Northeastern Naturalist S.R. Smedley and T. Wickman 2017 ii Vol. 24, Special Issue 7 Winter Ecology: Insights from Biology and History Preface Scott R. Smedley1 and Thomas Wickman2 All life as we know it, including our own species, inhabits a planet that rotates on a tilted axis. Consequently, winter is a reality for organisms that are denizens of the higher latitudes during that period in the Earth’s annual orbit when these regions receive limited solar input due to this tilt. For some species, such as subnivean inhabitants or flightless insects that disperse across the snow surface, winter brings benefits. However, it is also a season of scarcity and physical extremes, shaping both the ecology of northeastern North America and also the manner in which humans have engaged with the area’s environments. We here introduce this Northeastern Naturalist special issue, a multidisciplinary endeavor to investigate winter in this region, through lenses of both ecology and environmental history. Over several decades, winter ecology has coalesced as a discipline. Studies with a wintertime focus were previously, and often remain today, scattered through the ecological literature. However, Peter Marchand’s (1987) Life in the Cold drew upon a variety of these disparate studies and focused them into an introduction to winter ecology. The field was further popularized with the publication of Bernd Heinrich’s (2003) Winter World, which in recounting the author’s own work and that of others, provided a more general audience with an engaging portrayal of organisms’ strategies for coping with winter environments. In their review of winter ecology in northeastern North America, Campbell et al. (2005) stressed progress in understanding how biogeochemical processes function in winter and encouraged ecologists to focus further attention to wintertime phenomena, particularly given that a more thorough awareness is required to evaluate and predict organismal and ecosystem responses to climate change. More recent studies of winter ecology have built upon earlier, often more physiologically based work, transferring that knowledge to manipulated environments within the field. For example, earlier understanding (Schmid 1982, Storey and Storey 1984) of the remarkable biochemical basis of freeze tolerance in Lithobates sylvaticus (LeConte) (Wood Frog) was recently drawn upon to examine how snow cover and the frogs’ late-fall selection of hibernation sites affect their overwintering survival (O’Connor and Rittenhouse 2016). Our grasp of marine winter ecology has substantially expanded in recent years. Diel vertical movements of zooplankton within the water column likely represent the largest daily natural migration of biomass on the planet and are known to be 1 Department of Biology, Trinity College, Hartford, CT 06106; scott.smedley@trincoll.edu. 2History Department, Program in American Studies, Trinity College, Hartford, CT 06106; thomas.wickman@trincoll.edu. Winter Ecology: Insights from Biology and History 2017 Northeastern Naturalist 24(Special Issue 7):ii–viii Northeastern Naturalist iii S.R. Smedley and T. Wickman 2017 Vol. 24, Special Issue 7 driven by cues from solar illumination (Brierley 2014). Interestingly, these diel migrations also take place in marine waters during the darkness of the Arctic winter. Last et al. (2016) have now shown that these daily wintertime migrations in the Arctic are due to moonlight inducing downward movement. The field of marine winter ecology has benefited tremendously from technological advances. Genomic analysis of environmental DNA has shown that marine bacterial diversity peaks in the winter at high latitudes (Ladau et al. 2013). Use of global location-sensing loggers on seabirds has revealed previously unknown information on their winter distribution far from shore (Tranquilla et al. 2013). Recent analyses based on data collected through citizen science have demonstrated a major shift in the winter range of migratory Aechmophorus spp. (grebes), potentially due to a change in the abundance of marine fish, which are the primary prey of these birds (Wilson et al. 2013). The interaction of winter ecology and climate change will likely negatively impact many members of the biota of northeastern North America. An impressive example of winter ecology involves the seasonal change in the coat color of the Lepus americanus Erxleben (Snowshoe Hare). Near the southern limit of its distribution, the Snowshoe Hare in Pennsylvania has undergone a reduction in range over the past 3 decades and is now restricted to more northerly portions of the state (Diefenbach et al. 2016). Other researchers have shown that decreased duration of snow cover has resulted in lower survival stemming from longer periods of camouflage mismatch of white hares against a dark background (Mills et al. 2013, Zimova et al. 2016). Winter ecology even permeates the popular press and other media. Over the past month, news reports have focused public attention on the extremely high mortality (70%) of Alces alces (L.) (Moose) calves in New England by the end of their first winter (Bidgood 2017). This loss appears to be due, in large part, to dramatic increases in the populations of Dermacentor albipictus (Packard) (Winter Tick). Infested in the fall, individual Moose often host thousands of these ectoparasites through the winter. Calves are frequently incapable of surviving the chronic blood loss to the ticks. A variety of factors, including warmer winter temperatures, seem linked to this increase in tick numbers. Also in the news is the National Park Service decision to reintroduce Canis lupus L. (Gray Wolf) to the Isle Royale wilderness ecosystem in Lake Superior (Mlot 2016). A natural influx of these predators to the island took place over a bridge of lake ice in the 1940s, setting the stage for nearly 6 decades of study of predator–prey interactions between the Wolves and Moose. However, the Wolf population has plummeted over the past decade, presumably due to a high incidence of inbreeding. Input of genetic diversity into the population through natural dispersal of Wolves to Isle Royale seems unlikely given that substantial lake ice formation is now a rare event. Inspired by this scientific field, scholars in the humanities and social sciences have begun to investigate humans’ changing relationships with winter environments. Until recently, historians of North America often portrayed winter as a “slack” time between growing seasons (Demos 2004:11), unworthy of serious analysis. Economists, anthropologists, and historians have studied some winterNortheastern Naturalist S.R. Smedley and T. Wickman 2017 iv Vol. 24, Special Issue 7 specific human activities, such as hunting and trapping (Carlos and Lewis 2010, Krech 1999), logging (Cox 2010), and skiing (Coleman 2004a), but with principal attention given to commercial imperatives or social contexts. Some of the best new work has a geographical focus on northwestern North America (Cruikshank 2005, Di Stefano 2013) and on the Canadian subarctic and Arctic regions (Adcock 2016, Coates and Morrison 2001). New scholarship on northeastern North America, including in this special issue, establishes a fuller understanding of how nature has made humans’ diverse winter activities possible and how intensive human management has affected winter landscapes over time. A greater focus on winter-active species has revealed new answers to key questions within the field of North American environmental history. English and French settler societies of northern North America, for example, found that long winters made it difficult to replicate their European-style agricultural landscapes, and the processes of ecological imperialism proceeded in halting and uneven fashion in wintertime (Crosby 1986). The Canadian North has been paradigmatic in this sense (Piper and Sandlos 2007), but what about the contingent processes as they played out at lower latitudes of the Northeast? The introduction of livestock proved challenging. In some places, pigs and cattle roamed the winter woods and foraged along shore, altering the balance of nature wherever they went, and in other places, hogs or horses suffered where deep snow slowed them down or covered up potential food sources (Anderson 2004, Cronon 1983, Mancall 2010), in contrast to indigenous ungulates such as Moose, which fared better in winter (W ickman 2015). The story of canids is similarly complicated. The systematic eradication of Wolves and other creatures deemed pests was often carried out in the Northeast during winter and early spring, when farmers could dedicate more time to killing predators (Coleman 2004b:88, Kawashima and Tone 1983). Dog populations surged, meanwhile. The winter ramifications of human–canid relationships for game animals or competing scavengers around human settlements are now being integrated into the larger narratives of changes in the land. An emerging interdisciplinary subfield on coevolution promises to stimulate further study (Russell 2011). Native Americans’ knowledge of the overwintering strategies of a wide range of plants and animals have given them enduring sources of resilience as they have coped with winter stresses associated with colonialism. The participation of Native Americans in the over-harvesting of Castor canadensis Kuhl (North American Beaver) and other furbearers has been documented and debated (Krech 1999), but a wider repertoire of winter subsistence practices and cultural knowledge seldom has been recognized by scholars (Mancall 2009, 2013). Interdisciplinary methodologies and community-based collaborations promise to reveal the depth and subtlety of place-based knowledge over time. Climate fluctuations have influenced humans’ relationships with winter environments for centuries. Lower temperatures during the most-extreme phases of the Little Ice Age (ca. 1300–1850) made cold-weather knowledge especially valuable to societies across the continent (Fagan 2000, Kupperman 1984, Wickman 2015). Americans’ earliest written observations about natural history and seasonal weather Northeastern Naturalist v S.R. Smedley and T. Wickman 2017 Vol. 24, Special Issue 7 patterns took place in the context of a changing climate (Fuller 1998, Zilberstein 2016). Henry David Thoreau (1888, 2008) devoted considerable attention to wintertime processes, and recent interdisciplinary work has used some of Thoreau’s records as baselines to index anthropogenic changes in the climate (Primack 2014). Numerous subfields in environmental history have yet to be “winterized”. Despite an explosion of urban environmental histories, few studies give sustained attention to northeastern cities’ artificial winter habitats, whether due to snow removal or fuel burning (McKelvey 1995, Mergen 1997, Meyer 2009). Rivers have received booklength studies, with nuanced attention to industrial pollution, engineering methods, and social conflicts, but the icing up of northern rivers could be historicized further in terms of its cultural, economic, and ecological significance. Similarly, as marine environmental history has come into its own, historians are just beginning to comprehend the unique dynamics of the sea in winter (Bolster 2012). And where history of science meets environmental history, scholars should chronicle and interpret the discipline of winter ecology itself, as it has been practiced in the field, developed in laboratories, and represented in classrooms. The general public in the Northeast typically responds to mild or severe winters with both biological and historical questions. During the severe snowstorms of 2014–2015, bloggers and journalists expressed concern for wildlife (Oder 2015) and used data from past winters as a yardstick to judge the abnormality of snow depths (Epstein 2015, Holthaus 2015, Lipman 2015). Mild winter weather garners fewer headlines but has raised concern for activists, from Sheila Watt-Cloutier (2015), who has written about a “right to be cold”, to snow-sport athletes who founded the climate-advocacy group Protect Our Winters (Protect our Winters 2017). Scholars working in multidisciplinary or interdisciplinary collaborations have an opportunity to respond to this public demand for nuanced perspective about the past, present, and future of life in the cold. Attitudes toward winter have long been integral to national identity in Canada and other northern countries (Adcock 2016, Coates and Morrison 2001), but in the 21st century, snow and ice have become central to planetary thinking and transnational collaboration. Specialized research is as needed as ever, but so is the big picture, and we hope this special issue provides both. Literature Cited Adcock, T. 2016. A Cold Kingdom. The Otter, 17 March 2016. Available online at http:// niche-canada.org/2016/03/17/a-cold-kingdom/. Accessed 21 February 2017. Anderson, V. 2004. Creatures of Empire: How Domestic Animals Transformed Early America. Oxford University Press, New York, NY. 322 pp. Bidgood, J. 2017. Ticks, thriving in warm weather, take a ghastly toll on New England Moose. The New York Times, 19 January 2017. Available online at https://nyti. ms/2k42k7q. Accessed 21 February 2017. Bolster, J. 2012. The Mortal Sea: Fishing the Atlantic in the Age of Sail. 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