Isle Royale National Park Research Articles

Exploring the Origins of Red Foxes (Vulpes vulpes) on Isle Royale

Isle Royale National Park is generally considered a pristine ecosystem, but the island archipelago has a long history of human impacts that have altered the island's mammal communities through extirpations and introductions. The origin of red foxes (Vulpes vulpes) on the islands are of particular interest given their ubiquity and uncertain colonization history. Red foxes were first reported on Isle Royale in 1925, shortly after the foundation of a small fox farm that began on Isle Royale in 1922. We sequenced two mitochondrial haplotypes from red fox scats collected on Isle Royale and compared them to haplotypes from the mainland surrounding Lake Superior, the predominant source of the island's other native mammals. Some Isle Royale foxes matched widespread haplotypes commonly found across Canada, but over half of our samples matched haplotypes previously detected only in Newfoundland. While we cannot conclude a singular origin, we offer a working hypothesis red foxes on Isle Royale are derived from a combination of natural colonization and human introduction. Specifically, we propose native red foxes may be admixed with fur-farmed foxes from an introduction in the early 20th century.

Emigration and First-Year Movements of Initial Wolf Translocations to Isle Royale

Canis lupis (Gray Wolf or Wolf) can move great distances with important consequences to the persistence of isolated populations. We used global position system (GPS) tracking technology to summarize the movements of 4 Wolves translocated to Isle Royale National Park, MI, in 2018–2019. We quantified aspects of movement behavior of individual Wolves following translocation to, and an emigration event from the Isle Royale ecosystem.

Skeletal and Dental Development Preserve Evidence of Energetic Stress in the Moose of Isle Royale

Food shortages can leave diagnostic, and in the case of the dentition, irreversible changes in mineralized tissue that persist into historical and fossil records. Consequently, developmental defects of tooth enamel might be used to track ungulate population irruptions or declines in resource availability but dental tissue’s capacity for preserving historical population density changes has yet to be investigated in wild populations. We test the ability of macroscopic enamel defects, mandible and metapodial lengths to track changes in the well-known insular moose population of Isle Royale National Park. Our study demonstrates that 1) a moose density threshold exists on the island above which there is a significant decrease in mandible and metatarsus length and a concomitant increase in enamel hypoplasias, 2) food limitation has a more pronounced effect on male than female skeletal and dental growth, and 3) combined data from tooth enamel hypoplasias and bone lengths reflect the relative density of this ungulate population, and should be broadly applicable to other ungulate osteological samples. Developmental defects in dental enamel were among the highest recorded in a wild population, and even during low-density intervals the population density of Isle Royale moose has been high enough to negatively impact skeletal and dental growth, indicating the comparatively poor health of this isolated century-old ecosystem.

Climate Change Is Likely to Alter Future Wolf – Moose – Forest Interactions at Isle Royale National Park, United States

We evaluated how climate change and variable rates of moose browsing intensity, as they relate to wolf predation, might impact the forests of Isle Royale National Park, MI USA by conducting a simulation modelling experiment. The experiment consisted of contrasting three different scenarios of wolf management and with a static (current conditions) and changing climate (high emissions). Our results suggest that the interactive effects of wolf predation and climate change are likely to be temporally variable and dependent on biogeographic and forest successional processes. During the first 50 years of simulations, when the effects of climate change were minor, higher simulated rates of predation by wolves reduced moose population densities, resulting in greater forest biomass and higher carrying capacities for moose. However, over the longer term, early successional and highly palatable aspen and birch forests transitioned to late successional spruce and fir forests, regardless of climate or predation intensity. After 50 years, the effects of climate change and predation were driven by effects on balsam fir, a late successional species that is highly palatable to moose. High-intensity predation of moose allowed balsam fir to persist over the long term but only under the current climate scenario. Climate change caused a reduction in balsam fir and the other boreal species that moose currently feed on, and the few temperate species found on this isolated island were unable to compensate for such reductions, causing strong declines in total forest biomass. The direct effects of moose population management via reintroduction of wolves may become increasingly ineffective as the climate continues to warm because the productivity of boreal plant species may not be sufficient to support a moose population at all, and the isolation of the island from mainland temperate tree species may reduce the likelihood of compensatory species migrations.

Aquatic Macrophytes are Seasonally Important Dietary Resources for Moose

Moose (Alces alces) are generalist herbivores, but are important aquatic-terrestrial ecotone specialists. Aquatic macrophytes are a high-quality food source for moose during summer, but the importance of aquatic food sources to the moose diet is difficult to study. We used stable isotope analysis of carbon and nitrogen from moose hooves and forage (terrestrial plants, aquatic macrophytes, and arboreal lichen) to assess the diet of moose at Isle Royale National Park, Michigan, USA, using Bayesian mixing models. We also evaluated the isotopic variability along chronologies of serially sampled hooves. Overall, our mixing models indicate that 13%–27% of the summer moose diet was aquatic in origin. Among moose that died during winter, body condition was impaired and hoof 15N was higher where aquatic habitats were sparse. Although isotope chronologies preserved in hooves could significantly enhance our understanding of ungulate foraging ecology, interpretation of such chronologies is presently limited by our lack of knowledge pertaining to hoof growth rate and seasonal growth variability related to age and health. Distinct isotopic values among terrestrial plants, aquatic macrophytes, and arboreal lichens indicate that continued methodological advances in stable isotope ecology will lead to more precise estimates of the contribution of aquatic feeding to moose population dynamics and other ungulates.

Negative frequency-dependent foraging behaviour in a generalist herbivore (Alces alces) and its stabilizing influence on food web dynamics.

Resource selection is widely appreciated to be context-dependent and shaped by both biological and abiotic factors. However, few studies have empirically assessed the extent to which selective foraging behaviour is dynamic and varies in response to environmental conditions for free-ranging animal populations. Here, we assessed the extent that forage selection fluctuated in response to different environmental conditions for a free-ranging herbivore, moose (Alces alces), in Isle Royale National Park, over a 10-year period. More precisely, we assessed how moose selection for coniferous versus deciduous forage in winter varied between geographic regions and in relation to (a) the relative frequency of forage types in the environment (e.g. frequency-dependent foraging behaviour), (b) moose abundance, (c) predation rate (by grey wolves) and (d) snow depth. These factors are potentially important for their influence on the energetics of foraging. We also built a series of food-chain models to assess the influence of dynamic foraging strategies on the stability of food webs. Our analysis indicates that moose exhibited negative frequency dependence, by selectively exploiting rare resources. Frequency-dependent foraging was further mediated by density-dependent processes, which are likely to be predation, moose abundance or some combination of both. In particular, frequency dependence was weaker in years when predation risk was high (i.e. when the ratio of moose to wolves was relatively low). Selection for conifers was also slightly weaker during deep snow years. The food-chain analysis indicates that the type of frequency-dependent foraging strategy exhibited by herbivores had important consequences for the stability of ecological communities. In particular, the dynamic foraging strategy that we observed in the empirical analysis (i.e. negative frequency dependence being mediated by density-dependent processes) was associated with more stable food web dynamics compared to fixed foraging strategies. The results of this study indicated that forage selection is a complex ecological process, varying in response to both biological (predation and moose density) and abiotic factors (snow depth) and over relatively small spatial scales (between regions). This study also provides a useful framework for assessing the influence of other aspects of foraging behaviour on the stability of food web dynamics.

Detecting dynamic system regime boundaries with Fisher information: the case of ecosystems

The direct measurement of the resilience (resistance to disturbances) of an ecosystem’s current regime (or “alternative stable state”) remains a key concern for managing human impacts on these ecosystems and their risk of collapse. Approaches which utilize statistics or information theory have demonstrated utility in identifying regime boundaries. Here, we use Fisher information to establish the limits of the resilience of a dynamic regime of a predator–prey system. This is important because previous studies using Fisher information focused on detecting whether a regime change has occurred, whereas here we are interested in determining how much an ecological system can vary its properties without a regime change occurring. We illustrate the theory with simple two species systems. We apply it first to a predator–prey model and then to a 60-year wolf–moose population dataset from Isle Royale National Park in Michigan, USA. We assess the resilience boundaries and the operating range of a system’s parameters without a regime change from entirely new criteria for Fisher information, oriented toward regime stability. The approach allows us to use system measurements to determine the shape and depth of the “cup” as defined by the broader resilience concept.Graphic abstractThe direct measurement of the resilience (resistance to disturbances) of an ecosystem’s current regime remains a key concern for managing human impacts on these ecosystems and their risk of collapse. Here, we use Fisher information to establish the limits of stability of a dynamic regime of a predator–prey system. The region of stability is represented by the “floor of the canyon” in the adjacent graphic. While the theory is illustrated with an ecosystem example, it is applicable in its present form to dynamic systems in general.

Response of boreal lakes to changing wind strength: Coherent physical changes across two large lakes but varying effects on primary producers over the 20th century

AbstractNear‐surface wind speeds have changed over recent decades, raising questions about the extent to which these changes are altering the vertical thermal structure of lakes and affecting lake food webs. Neo‐ and paleolimnological techniques were used to assess wind‐driven changes in lake thermal habitat and resulting effects on primary producers in two lakes in Isle Royale National Park, an island archipelago located in Lake Superior, where wind speed has increased in recent decades. Responses in Siskiwit Lake, a large (16 km2 surface area), deep (Zmax = 49 m), oligotrophic lake, were compared to those of Lake Desor, a moderately large (4.3 km2) but shallower (Zmax = 13 m), mesotrophic lake. High‐frequency sensor data suggested that changes in wind speed affected epilimnion thickness in both lakes synchronously (ρ = 0.7, p < 0.001). Diatom‐inferred mixing depths suggested a coherent shift in both lakes to deeper mixing (an increase of 3 and 6 m) since 1920 (ρ = 0.8), which was correlated with an increase in regional wind speed during the 20th century at the decadal‐scale in Lake Desor and Siskiwit Lake (ρ = 0.6 and 0.4, respectively). In Lake Desor, algal biomass declined as mixing deepened from 1920 to 1980, and then cyanobacteria and cryptophyte pigments increased from 1980 to present, a period of inferred stable and deep mixing. Algal pigment concentrations in Siskiwit Lake were unchanged as mixing depth deepened. Although changes in wind speed altered lake physical structure similarly, the ecological consequences of these changes differed between lakes and were most likely influenced by lake‐specific variability in nutrient and light availability.

The relative roles of dispersal and establishment for shaping whole-lake aquatic macrophyte richness, diversity, and community structure in hemiboreal lakes

ABSTRACTAquatic macrophyte diversity and structure play an important role in freshwater communities, yet little research on macrophytes has focused on native communities in the undisturbed hemiboreal lakes of North America. We used data collected from 15 inland lakes on Isle Royale National Park (Michigan, USA) to identify relationships among macrophyte metrics that describe dispersal and introduction potential (richness, diversity, community structure via growth form, and connectivity) and lake metrics that control establishment and growth potential. Results from partial least square regression analyses found that the most important and recurring predictors were those corresponding to establishment and growth. Alkalinity was the major contributor to the variation seen in richness, Shannon evenness, and Simpson inverse diversity metrics. Sediment phosphorus also contributed significantly to richness and diversity, but negatively. For growth form, half of the variation in submersed species related to alkalinity while almost half of variation in emergent species related to alkalinity and water color. Most of the variation in floating-leaved species was negatively related to alkalinity and water color. Our results suggest a high relative importance of establishment and growth metrics compared to introduction and dispersal metrics for understanding native macrophyte communities in North American hemiboreal lakes.

Dynamic colonization history in a rediscovered Isle Royale carnivore

Island ecosystems are globally threatened, and efforts to restore historical communities are widespread. Such conservation efforts should be informed by accurate assessments of historical community composition to establish appropriate restoration targets. Isle Royale National Park is one of the most researched island ecosystems in the world, yet little is actually known about the biogeographic history of most Isle Royale taxa. To address this uncertainty and inform restoration targets, we determined the phylogeographic history of American martens (Martes americana), a species rediscovered on Isle Royale 76 years after presumed extirpation. We characterized the genetic composition of martens throughout the Great Lakes region using nuclear and mitochondrial markers, identified the source of Isle Royale martens using genetic structure analyses, and used demographic bottleneck tests to evaluate (eliminate redundancy of test). 3 competing colonization scenarios. Martens exhibited significant structure regionally, including a distinct Isle Royale cluster, but mitochondrial sequences revealed no monophyletic clades or evolutionarily significant units. Rather, martens were historically extirpated and recolonized Isle Royale from neighbouring Ontario, Canada in the late 20th century. These findings illustrate the underappreciated dynamics of island communities, underscore the importance of historical biogeography for establishing restoration baselines, and provide optimism for extirpated and declining Isle Royale vertebrates whose reintroductions have been widely debated.

Effects of Climate Change on Lake Thermal Structure and Biotic Response in Northern Wilderness Lakes

Climate disrupts aquatic ecosystems directly through changes in temperature, wind, and precipitation, and indirectly through watershed effects. Climate-induced changes in northern lakes include longer ice-free season, stronger stratification, browning, shifts in algae, and more cyanobacterial blooms. We compared retrospective temperature-depth relationships modeled using MINLAKE2012 with biogeochemical changes recorded in sediment cores. Four lakes in Voyageurs National Park (VOYA) and four lakes in Isle Royale National Park (ISRO) were studied. Meteorological data from International Falls and Duluth, Minnesota, were used for VOYA and ISRO, respectively. Model output was processed to analyze epilimnetic and hypolimnetic water temperatures and thermal gradients between two periods (1962–1986, 1987–2011). Common trends were increased summer epilimnion temperatures and, for deep lakes, increased frequency and duration of thermoclines. Changes in diatom communities differed between shallow and deep lakes and the parks. Based on changes in benthic and tychoplanktonic communities, shallow lake diatoms respond to temperature, mixing events, pH, and habitat. Changes in deep lakes are evident in the deep chlorophyll layer community of Cyclotella and Discostella species, mirroring modeled changes in thermocline depth and stability, and in Asterionella and Fragilaria species, reflecting the indirect effects of in-lake and watershed nutrient cycling and spring mixing.

Modelling moose-forest interactions under different predation scenarios at Isle Royale National Park, USA.

Loss of top predators may contribute to high ungulate population densities and chronic over-browsing of forest ecosystems. However, spatial and temporal variability in the strength of interactions between predators and ungulates occurs over scales that are much shorter than the scales over which forest communities change, making it difficult to characterize trophic cascades in forest ecosystems. We applied the LANDIS-II forest succession model and a recently developed ungulate browsing extension to model how the moose population could interact with the forest ecosystem of Isle Royale National Park, USA, under three different wolf predation scenarios. We contrasted a 100-yr future without wolves (no predation) with two predation scenarios (weak, long-term average predation rates and strong,higher than average rates). Increasing predation rates led to lower peak moose population densities, lower biomass removal rates, and higher estimates of forage availability and landscape carrying capacity, especially during the first 40 yr of simulations. Thereafter, moose population density was similar for all predation scenarios, but available forage biomass and the carrying capacity of the landscape continued to diverge among predation scenarios. Changes in total aboveground live biomass and species composition were most pronounced in the no predation and weak predation scenarios. Consistent with smaller-scale studies, high browsing rates led to reductions in the biomass of heavily browsed Populus tremuloides, Betula papyrifera, and Abies balsamea, and increases in the biomass of unbrowsed Picea glauca and Piceamariana, especially after the simulation year 2050, when existing boreal hardwood stands at Isle Royale are projected to senesce. As a consequence, lower predation rates corresponded with a landscape that progressively shifted toward dominance by Piceaglauca and Piceamariana, and lacking available forage biomass. Consistencies with previously documented small-scale successional shifts, and population estimates and trends that approximate those from this and other boreal forests that support moose provide some confidence that these dynamics represent a trophic cascade and therefore provide an important baseline against which to evaluate long-term and large-scale effects of alternative predator management strategies on ungulate populations and forest succession.

Modeling Viability of a Potential Canada Lynx Reintroduction to Isle Royale National Park

Canada lynx (Lynx canadensis) were extirpated from Isle Royale in the 1930s. We conducted a population viability analysis (PVA) of a potential lynx reintroduction to better understand viability, uncertainty, and management options. We estimated that the 544-km2 island can support 30 lynx. The probability of 100-year population persistence was 0.36 for a model that simulated a decadal lynx-hare population cycle. A noncyclic model predicted a 0.73 probability of 100-year persistence. Inbreeding depression had a substantial negative effect on modeled persistence. Historically, periodic immigration of mainland lynx via an ice bridge probably reduced or prevented inbreeding depression on the island. The introduction of one male and one female lynx every 10 years increased the probability of 100-year persistence to 0.98 in the cyclic model. Occasional anthropogenic transfers of lynx to the island might be necessary because the frequency of ice bridge formation has decreased. However, our baseline models might underestimate viability because they used demographic rates from mainland studies where lynx were exposed to anthropogenic mortality. When we removed assumed anthropogenic mortality—which should be negligible on Isle Royale—the probability of 100-year persistence was 0.99 for the cyclic model, even without supplementation. Reintroducing lynx to Isle Royale appears feasible, assuming appropriate monitoring and management. Reintroducing lynx would restore a missing native species to Isle Royale and would increase our understanding of lynx ecology.

Simulating ungulate herbivory across forest landscapes: A browsing extension for LANDIS-II

Browsing ungulates alter forest productivity and vegetation succession through selective foraging on species that often dominate early succession. However, the long-term and large-scale effects of browsing on forest succession are not possible to project without the use of simulation models. To explore the effects of ungulates on succession in a spatially explicit manner, we developed a Browse Extension that simulates the effects of browsing ungulates on the growth and survival of plant species cohorts within the LANDIS-II spatially dynamic forest landscape simulation model framework. We demonstrate the capabilities of the new extension and explore the spatial effects of ungulates on forest composition and dynamics using two case studies. The first case study examined the long-term effects of persistently high white-tailed deer browsing rates in the northern hardwood forests of the Allegheny National Forest, USA. In the second case study, we incorporated a dynamic ungulate population model to simulate interactions between the moose population and boreal forest landscape of Isle Royale National Park, USA. In both model applications, browsing reduced total aboveground live biomass and caused shifts in forest composition. Simulations that included effects of browsing resulted in successional patterns that were more similar to those observed in the study regions compared to simulations that did not incorporate browsing effects. Further, model estimates of moose population density and available forage biomass were similar to previously published field estimates at Isle Royale and in other moose-boreal forest systems. Our simulations suggest that neglecting effects of browsing when modeling forest succession in ecosystems known to be influenced by ungulates may result in flawed predictions of aboveground biomass and tree species composition.

Reproduction of Lake Trout Morphotypes at Isle Royale in Northern Lake Superior

AbstractThe reproductive biology of male and female lean, siscowet, redfin, and humper Lake Trout Salvelinus namaycush was assessed by measuring the gonadosomatic index (GSI) and by histological analysis of the gonads over 2 years from spring to fall at sites surrounding Isle Royale National Park, Lake Superior. Whole livers, otoliths, and ovaries were also collected to measure hepatosomatic index (HSI), age, and fecundity. In general, GSIs increased from spring to summer in male and female siscowets, leans, redfins, and humpers, and this was accompanied by a large increase in the number of vitellogenic (>2.0 mm) follicles in females. Postspawned ovaries and regressing testes were predominant in all morphotypes by October but were also observed in September, indicating that some spawning occurred at least by September. Further, preovulatory siscowet females (GSIs > 10) and siscowet males with fully mature and even regressing testes were observed in the spring and summer, indicating that spawning occurs at least in the spring and fall in siscowets at Isle Royale. For all morphotypes, average male HSIs were significantly lower than female HSIs. Absolute fecundity was positively related to size and age for all morphotypes. Humpers had the lowest absolute fecundity. Leans, redfins, and siscowets had similar fecundity relationships. The results conclusively demonstrate that deepwater siscowet Lake Trout reproduce at least during the spring and fall at Isle Royale and that leans, humpers, and redfins reproduce during the fall. In lakes that exhibit sympatric populations of deepwater and shallow‐water ecotypes of Arctic Char Salvelinus alpinus, the deepwater form also spawns in both spring and fall. Our data and those on Arctic Char suggest that there is an adaptive strategy for maintaining alternative spawning times in the deepwater ecotypes in these populations.Received June 20, 2016; accepted November 2, 2016 Published online January 31, 2017

A regional assessment of chemicals of concern in surface waters of four Midwestern United States national parks.

Anthropogenic chemicals and their potential for adverse biological effects raise concern for aquatic ecosystem health in protected areas. During 2013-15, surface waters of four Midwestern United States national parks were sampled and analyzed for wastewater indicators, pharmaceuticals, personal care products, and pesticides. More chemicals and higher concentrations were detected at the two parks with greater urban influences (Mississippi National River and Recreation Area and Indiana Dunes National Lakeshore) than at the two more remote parks (Apostle Islands National Lakeshore and Isle Royale National Park). Atrazine (10-15ng/L) and N,N-diethyl-meta-toluamide (16-120ng/L) were the only chemicals detected in inland lakes of a remote island national park (Isle Royale National Park). Bisphenol A and organophosphate flame retardants were commonly detected at the other sampled parks. Gabapentin and simazine had the highest observed concentrations (>1000ng/L) in three and two samples, respectively. At the two parks with urban influences, metolachlor and simazine concentrations were similar to those reported for other major urban rivers in the United States. Environmental concentrations of detected chemicals were often orders of magnitude less than standards or reference values with three exceptions: (1) hydrochlorothiazide exceeded a human health-based screening value in seven samples, (2) estrone exceeded a predicted critical environmental concentration for fish pharmacological effects in one sample, and (3) simazine was approaching the 4000ng/L Maximum Contaminant Level in one sample even though this concentration is not expected to reflect peak pesticide use. Although few environmental concentrations were approaching or exceeded standards or reference values, concentrations were often in ranges reported to elicit effects in aquatic biota. Data from this study will assist in establishing a baseline for chemicals of concern in Midwestern national parks and highlight the need to better understand the sources, pathways, and potential adverse effects to aquatic systems in national parks.

Wolf Listeners: An Introduction to the Acoustemological Politics and Poetics of Isle Royale National Park

Listening to wolf howls as both material object and socially constructed metaphor highlights the contested relationship between nature and culture. The author conducted field research on Isle Royale National Park from 2011 to 2015, from which data he offers a narrative wherein citizen-scientists who listen for the howl literally “lend their ears” to a wolf biologist who has led the longest continuous predator-prey study in the world. The theoretical framework of this essay extends acoustic ecology, first theorized by R. Murray Schafer, to include environmental history and cultural theory, which problematizes definitions of “nature” and “natural.” Ultimately, this introduction describes a nuanced form of participatory, situational environmental music that plays out in the everyday lives of those listening on this remote, roadless island on Lake Superior.

Mayflies, stoneflies, and caddisflies of streams and marshes of Indiana Dunes National Lakeshore, USA.

United States National Parks have protected natural communities for one hundred years. Indiana Dunes National Lakeshore (INDU) is a park unit along the southern boundary of Lake Michigan in Indiana, USA. An inventory of 19 sites, consisting of a seep, 12 streams, four marshes, a bog, and a fen were examined for mayflies (Ephemeroptera), stoneflies (Plecoptera), and caddisflies (Trichoptera) (EPT taxa). Volunteers and authors collect 35 ultraviolet light traps during summer 2013 and supplementary benthic and adult sampling added species not attracted by lights or that were only present in colder months. Seventy-eight EPT species were recovered: 12 mayflies, two stoneflies, and 64 caddisflies. The EPT richness found at INDU was a low proportion of the number of species known from Indiana: caddisflies contributed only 32.7% of known state fauna, mayflies and stoneflies contributed 8.4% and 2.3%, respectively. Site EPT richness ranged from one for a seep to 34 for an 8 m-wide stream. Richness in streams generally increased with stream size. Seven new state records and rare species are reported. The number of EPT species at INDU is slightly larger than that found at Isle Royale National Park in 2013, and the community composition and evenness between orders were different.

Ephemeroptera, Plecoptera, and Trichoptera on Isle Royale National Park, USA, compared to mainland species pool and size distribution.

Extensive sampling for aquatic insects was conducted in the orders Ephemeroptera (mayflies), Plecoptera (stoneflies), and Trichoptera (caddisflies) (EPT) of Isle Royale National Park (ISRO), Michigan, United States of America, during summer 2013. The island was ice covered until 8,000 to 10,000 years ago and is isolated by 22–70 km distance from the mainland. Two hypotheses were examined: that ISRO EPT richness would be much reduced from the mainland, and that the species colonizing ISRO would be of smaller size than mainland, adults presumably using updrafts to bridge the distance from mainland sources. Data sets were developed for known mainland EPT species and size for those species. The first hypothesis was confirmed with the mainland species pool consisting of 417 EPT, while ISRO is known to support 73 species. Richness of EPT is directly related to the number of specimens examined. Small streams supported five EPT species, while 15–25 species were found in larger streams. Lakeshores had intermediate diversity. The second hypothesis was substantiated for stoneflies, but not for mayflies or caddisflies. Stoneflies apparently are poorer fliers than either of the other two orders.

Field philosophy: environmental learning and moral development in Isle Royale National Park

We conducted a qualitative analysis of 5 years of student writing data to understand learning and moral development on a field philosophy course in Isle Royale National Park. We were interested in the connection between physical experiences in the natural world and the way students care about or value nonhuman beings, natural systems, and place. Students consistently demonstrated a shift from a dualistic to a more complex understanding of ideas, relationships, and the natural world. This widening perception and moral awareness occurred in tandem with course content learning. In this paper we describe the model that emerged from our data to describe this shift from dualism to complexity – from self-awareness, to social learning, to curriculum engagement, to a wider moral community – and the factors that facilitate each step. We also provide a planning and assessment tool to help other educators begin to observe ethical development and relational growth in their own interdisciplinary environmental field classes.