Birds in Winter: Surviving the Most Challenging Season

Birds in Winter

Surviving the Most Challenging Season

Roger F. Pasquier

Illustrations by Margaret La Farge

Princeton University Press

Princeton and Oxford

Copyright © 2019 by Princeton University Press

Published by Princeton University Press

41 William Street, Princeton, New Jersey 08540

6 Oxford Street, Woodstock, Oxfordshire OX20 1TR

press.princeton.edu

All Rights Reserved

LCCN 2018958180

ISBN 978-069-117855-4

eISBN 978-069-119543-8 (ebook)

Version 1.0

British Library Cataloging-in-Publication Data is available

Editorial: Robert Kirk and Kristin Zodrow

Production Editorial: Ellen Foos

Text Design: D & N Publishing, Wiltshire, UK

Jacket Design: Lorraine Doneker

Jacket image: Markus Varesvuo/Agami Picture Library

Production: Steven Sears

Publicity: Matthew Taylor and Julia Hall

Copyeditor: Laurel Anderton

For

Phoebe Goodhue Milliken

CONTENTS

Introduction  7

Acknowledgments  10

Chapter 1:  Responding to Winter  11

Chapter 2:  Preparing for Winter  35

Chapter 3:  Winter Ranges and Habitat Selection in Migratory Birds  53

Chapter 4:  Spatial and Social Organization  81

Chapter 5:  Survival  121

Chapter 6:  The Winter Day  149

Chapter 7:  Anticipating Spring  167

Chapter 8:  Departure  187

Chapter 9:  Conservation  207

Chapter 10: Climate Change  233

Bibliography  251

Index  284

Male Black-billed Capercaillies in Siberia eat larch twigs through the winter, picking whatever is above the snow. This increases the number of shoots that will grow from the stem the next spring, providing food the following winter.

INTRODUCTION

IN 1976, WHEN I was writing Watching Birds: An Introduction to Ornithology, I had an abundance of books and papers in scientific journals to draw from as I described the breeding behavior and migrations of birds. Conspicuously missing, however, was a comparable level of information on the remaining portion of the annual cycle—winter—which to me seemed no less interesting, and no less vital to understanding the lives of birds. I believe the chapter on winter I wrote for Watching Birds was then the first unified, albeit modest, account of the ecology and behavior of how both permanent residents and migrants deal with that season. I was sure there was much more to be researched and brought together, and I immediately proposed the idea to my editor as my next book. Too narrow a topic! was the response then, as well as later in a succession of intervals, each of about fifteen years, when I proposed it again to subsequent editors.

Finally, in 2015, after completing Painting Central Park, another book I had long wanted to write, I decided to take the plunge, research the topic of birds in winter in all its aspects, and determine for myself whether there was a book in it. Over those nearly 40 intervening years, winter had also come to interest a new generation of ornithologists. The Smithsonian Institution’s 1977 and 1989 symposia on the ecology and conservation of migrant birds in the Neotropics, for example, brought together tremendous amounts of essential information, inspired even more research, and produced publications that are still primary references. The years around the turn of this century saw expanded interest in austral migration and winter in Australia and South America—where, in fact, W. H. Hudson had vividly depicted it in Far Away and Long Ago (1918) based on his boyhood memories of Argentina in the 1870s.

Over these decades, ornithologists in Finland and Scandinavia were publishing a steady stream of papers describing the social and ecological winter dynamics of boreal forest birds, while from Britain and the Netherlands were coming accounts of how shorebirds spend that season in northerly latitudes, permitting useful comparisons with the North American and Eurasian shorebirds that winter far south of the equator. Similarly, in North America intensive long-term studies of chickadees and juncos have revealed the complexities of their winter lives. (Sadly, there is still very little information about the birds wintering in Southeast Asia, where forests are being cut at an alarming rate, or from much of Africa, where more frequent and intense droughts are transforming the landscape.) The recent rapid advances in satellite tracking technology and stable isotope analysis have enabled scientists to follow individual birds day by day and practically meal by meal all through the winter when these birds are continents and oceans away. And today the growing concern about conservation, especially the loss of wintering habitat, and the impacts of climate change have added new perspectives and new urgency to understanding the entire annual cycle of birds.

So, in 2015 I was glad to return to a seat at the same table in the library of the Department of Ornithology in the American Museum of Natural History where I had written Watching Birds. The perfect place to research and write. I’ve never had more fun! When discussing the feathered toes of the Snowy Owl, the differences in bill lengths of North American and Caribbean kingbirds and vireos, or a species like the Tibetan Ground Tit that I did not know at all, I could look at the specimens. It was hard to say basta! to research when every newest issue of one of more than 20 ornithological, behavioral, ecological, and conservation journals I was following might have something amazing and irresistible to include. I drew the line at the last of the mid-2018 publications—and hope I will not soon regret it.

Of course, when I described my project, some people thought my topic was simply the familiar birds of northern latitudes that stay where it is cold. But no, winter is a global phenomenon, and it influences birds all over the world, including many of those that never leave the tropics but must share their habitat during parts of the year with birds from the Northern or Southern Hemisphere, or from both. We routinely say Blackburnian Warblers winter in Colombia and White Storks winter in Africa, so there seemed every reason to follow them there to learn the challenges of that season and how the birds fit into those ecosystems. (I have kept the discussion of how migrants reach their winter destinations to a minimum, since today there are more books on migration, dazzling in their comprehensiveness, than ever before.) And, since life is a continual cycle, what happens to birds during the winter affects their subsequent breeding season, often in unexpected ways, as new technology is revealing. Finally, since one of the greatest pleasures of watching birds is knowing how far some have traveled and how their physical and behavioral adaptations have enabled them to survive in what may be widely different environments, one surely wants to include the birds that winter where it may not seem like winter at all.

Another of the pleasures of watching birds is understanding more of what’s really going on among the most familiar species that we may see all year or through the winter when many birds are so much more visible. In the midlatitudes, where in winter there may be no concealing foliage, it is much easier to follow birds through the day. Many also seem tamer, perhaps because of hunger and, in extreme cold, their tendency to reduce movement to save energy. Naive young birds living through their first winter sometimes give us glimpses experienced adults do not allow. I have seen a first-year Red-tailed Hawk bathing in a stream and a young Northern Goshawk carelessly bold.

Showing readers how much there is to see through winter, when so many birders are least active, has been one of my goals. On my daily walks through Central Park to and from the museum as I was writing this book, I observed with new eyes some of what I was researching and writing—and smiled as I thought, "I know now what you’re doing." I saw woodpeckers, jays, and tits beginning to store food while it still seemed like summer, and interactions at feeders starting in November as different species made others give way and as individuals asserted their rank in a flock. The robins and grackles that used to be harbingers of spring in New York were now commonly foraging successfully on the lawns through our mostly snowless winters. As the males of each of the duck species that winters on the lakes and reservoir in the park completed their molt, I saw the beginnings of their courtship. By February, when most of them were paired, I looked with sympathy at the extra males that had had no luck. And also in February the titmice and cardinals began singing, weeks before either would actually start nesting.

I hope, therefore, readers will look with new insight, just as I did, at the behavior of the birds around them through the winter and will consider how many of the physical and ecological adaptations of all birds result from selective pressure during the season when survival is the priority wherever birds are. The consequences of molt and migration schedules, of rank due to age, gender, and relative body size, winter separations of competing populations and species by latitude and habitat, seasonal rainfall shifts, incidents of extreme weather, and other factors all interweave to make winter at any latitude more challenging for most birds than their breeding season. And, as the very nature of winter is now changing, from the drying of many tropical regions to the warming of polar ocean currents and the acceleration of spring events faster than some birds can shift to that season, we need to appreciate the scope of these impacts so we can, individually and collectively, take appropriate action to preserve and restore vital habitats and implement the practices that will reduce rates of climate change. Winter is too fascinating a season to lose!

ACKNOWLEDGMENTS

MY PRIMARY DEBT, of course, is to the legions of scientists who have worked through intense cold or extreme heat, unending nights, too short days, rolling seas, and long years to learn firsthand what birds are really doing during winter. Some of them I would have liked to join, especially those working in habitats that have since been degraded or lost. It is the diligence and insights of all these scientists that have provided me the material necessary to create this book. Two anonymous readers from that community made helpful suggestions on the manuscript, mainly about its structure. Any errors of fact or interpretation are my own.

In the Department of Ornithology at the American Museum, Joel Cracraft, Paul Sweet, Lydia Geratano, Peter Capainolo, and Mary LeCroy all literally and figuratively opened doors for me. In the museum’s main library, Tom Baione, Mai Reitmeyer, and Annette Springer showed me through the several floors of stacks and helped me find other publications available only on the internet.

At the American Bird Conservancy, Michael Parr, Aditi Desai, and Grant Sizemore, and my former colleagues at the National Audubon Society, Matt Jeffery and Gary Langham, gave me information on current conservation activities and examples of their organizations’ successes in protecting the habitats of wintering birds.

Margaret La Farge’s line drawings animate these pages. I wish there could have been more!

At the Princeton University Press, my editor Robert Kirk was supportive from the very beginning and especially helpful when I needed advice on reconfiguring some of the chapters. Ellen Foos and Kristin Zodrow meticulously oversaw all the steps that metamorphose a larval typescript into perhaps a butterfly of a book. Laurel Anderton, the copy editor, spotted words and numbers in the text needing correction with the acuity of a kinglet making its way through the winter woods, finding prey no other bird could see. Thanks to them all.

CHAPTER 1

RESPONDING TO WINTER

AS LONG AS EARTH has circled the sun and tilted on its axis during the course of an annual rotation, some parts of the planet have received less heat and light for a portion of each year. This annual tilting of each pole toward and then away from the sun produces the seasons, the coldest and darkest being what we call winter.

In some earlier epochs of Earth’s history, the entire planet was much warmer than it is today, with subtropical conditions extending as far as latitude 50° north, and the impacts of winter were probably minimal; at other times they were more intense.

Birds, on Earth 155 million or more years, have evolved and responded to the opportunities that these changing conditions have presented to survival and reproduction. The last five million years have been Earth’s coolest, and therefore most seasonal. During the last 1.8 million years there have been at least 22 major periods of glaciation that made the entire planet colder and drier. Most of the birds species alive today lived through at least some of these oscillations, and by the end of the most recent glaciation, about 12,000 years ago, probably all of them had evolved their current form and annual life cycle.

This annual cycle, once a bird is an adult, can most easily be divided into the time spent in reproductive activities—finding a mate, nesting, and raising young until their independence—and the rest of the year, when each bird fends for itself. For the birds that must deal with winter—at least a third of the species living today—this season usually runs longer than the breeding season and is more challenging. At the higher latitudes of all continents, north and south, the features of winter include lower ambient temperatures, increased wind, and fewer hours of daylight.

These factors directly affect many of the food sources that were consumed by birds during the warmth of the breeding season. Diminished sunlight and increased cold substantially reduce or curtail production of plant material, in turn diminishing the abundance or availability of insects and other small animals that feed on plants, while snow may cover the ground and many water bodies may freeze, further limiting accessibility of food. For what food remains, there are fewer hours with enough light to forage, and longer, colder nights during which birds must fast, burning the energy they have gained through what they consumed the previous day. For small birds, those weighing less than 100 grams, insulation against cold is another challenge.

Warm-bloodedness has prevented birds from simply shutting down through an annual cold period, as can reptiles and amphibians, while the mobility that comes with flight has enabled them to search more widely for the food and shelter they need to survive, whether over permanent home ranges or more distant hospitable latitudes. Birds’ responses to winter vary. Some stay in place, others migrate annually from and then back to the breeding site, while a smaller number are nomadic or have periodic irruptions when populations build beyond what the local winter food supply can sustain. Winter therefore affects not only the birds that remain to endure it and those that travel from higher latitudes to avoid it, but also the birds in regions where migrants settle, creating potential competition between residents and transients, often at a time when equatorial weather patterns also influence the availability of food.

Since migration is the response of most birds of high and middle latitudes, we will consider it first.

MIGRATION

ORIGINS OF MIGRATION

The fossil remains of birds show that some had the anatomy required for long-distance flight at least 100 million years ago. Climatic conditions have probably prompted migration in many current bird families for the past 15 million years or longer (Steadman 2005). Migration evolved among both birds of high latitudes and those of the tropics, where seasonal variations in rainfall and the resulting abundance or decline of food resources led to more local annual movements.

For birds originating at high latitudes, the eras of cold forced a retreat toward lower latitudes where exploitable habitat remained. Through the annual and long-term fluctuations in warmth, the birds were able to return poleward to breed. Their migrations to escape winter were relatively short and may not have taken place every year if birds could survive without moving. During glacial periods, the range of some birds contracted, but these birds had already evolved to exploit resources available during the cold season and were able to remain farther north than birds of tropical origin that may have been their summer neighbors.

In North America, most of the nonmigrant or short-distance landbird migrants are from families that originated in the Palearctic and crossed the land bridge that at various times of glacial maxima joined Siberia and Alaska. These include woodpeckers, corvids, tits, nuthatches, and finches, in which some species, or populations within species, are nonmigratory and others are short-distance migrants or have irregular movements in response to annual variations in food supply. For them, lifetime reproductive potential may be greater when they can remain on their breeding territory or have only a short migration to reach it; the risk is that they may not survive through the cold winter.

Long-distance migrants retreating to the tropics, in contrast, may increase their chance of survival by avoiding the cold, even if they reduce their reproductive potential the next spring by using time to establish a new territory and by missing some of the food sources the residents and earlier arrivals will already have consumed. This must, however, be an acceptable trade-off, since today, in all hemispheres, a large portion of the species breeding at higher latitudes migrate to the tropics. Some are from families that originated in the tropics, such as the uniquely Western Hemisphere hummingbirds and tyrant flycatchers. Other families that have many long-distance migrants—for example, wood warblers and blackbirds—are descended from birds that, like the ancestors of many families with short-distance migrants, came across Beringia from the Old World and then radiated into the families we know today in the Western Hemisphere. Parts of the radiation into current species may have taken place in the tropics. These families have been present long enough that some members could have evolved migratory behavior, lost it, and evolved it again several times in response to different climatic conditions. Thus, some of the migratory species living today may owe their habit to ancestors that originally retreated from cold weather at northern latitudes, while others are descended from birds that later expanded north out of the tropics (Winger et al. 2014). Since some of these families of Old World origin, such as sparrows, also include species that expanded far into the Southern Hemisphere and now migrate back toward tropical latitudes to escape the austral winter, both scenarios have been factors at different stages of their families’ presence in the New World. Similarly, among Palearctic birds that go to Africa in winter, nearly all have breeding populations or close relatives there that may have been the ancestral source of the current migrants (Bohning-Gaese 2005).

Why would birds in the tropics leave? Competition for resources between species, and for breeding space within species, may, in the interglacial eras when regions farther from the equator warmed, have led some birds, especially young ones seeking their first territories, to edge their way beyond their birthplace. In the same way, on a shorter timescale, changes in habitat and climate over the last century have enabled several North American species to expand their breeding range significantly northward, with some retreating to their historical range each winter. The seasonal blooms of food sources at higher latitudes are short but intense, providing brief abundance of protein-rich food ideal for feeding nestlings; these resources, however, diminish rapidly, forcing a retreat back to warmer regions. During the warmer interglacial periods, such as today, the distance between exploitable breeding habitat and viable wintering areas was longer than in the glacial phases when cold reduced how far tropical birds could spread from their place of origin. The breeding ranges of many birds expanding from the tropics therefore went through periods of growth and contraction, while the winter range to which they returned each year may have been more stable.

Many tropical birds have annual seasonal movements that make it a short evolutionary step to long-distance migration. The Yellow-green Vireo (Vireo flavoviridis), for example, breeds from northern Mexico to Panama; in Panama, most have left by the end of August for the upper Amazon basin, returning in December. Their diet is small fruits, and their migration is linked to their food’s annual cycle of abundance and decline (Morton 1977). Other intertropical migrants do not traverse significant latitudes; many, like the Resplendent Quetzal (Pharomachrus mocinno) of Mexico and Central America, move up and down mountains following the timetable of fruiting plants at different elevations. From this intratropical origin, it was an easy evolutionary step during interglacial periods for some birds within the tropics to expand their ranges north or south as changes in climate prompted the growth of food sources not previously available (Rappole and Tipton 1992).

The 1.8 million years of glacial and interglacial periods have given birds of both tropical and temperate origin time to evolve complex variations of these general patterns, refined by modern species in the last several thousand years when Earth’s climate has been most like today’s. The Western Hemisphere thrush genus Catharus, for example, contains 12 species; 7 are restricted to the Neotropics, where the genus originated, and 5 breed in North America: the Bicknell’s (C. bicknelli), Gray-cheeked (C. minimus), Hermit (C. guttatus), and Swainson’s (C. ustulatus) Thrushes and the Veery (C. fuscescens). All but the Hermit Thrush return entirely to various parts of the tropics in winter; the Hermit Thrush is a short-distance migrant wintering primarily from the central and southern United States to Mexico and Guatemala.

TYPES OF MIGRATION

Migration and other less regular large-scale movements to avoid winter take many forms. These reflect the origins of each species as well as its adaptations to different habitats over the course of a year.

PARTIAL MIGRATION Many birds have both migrant and resident populations. Some individuals move to lower latitudes, while others remain in the breeding range year round. In the Northern Hemisphere, the entire high-latitude population may shift southward into the permanent range of the southern population and beyond, or some individuals from all parts of the range may move while others remain. Often, adult males stay closest to the breeding site, the better to regain their territory for the following year, while females and young move farther (Gauthreaux 1982). In parts of the range where birds are found all year, it is impossible without marking every bird to determine which are residents and which are migrants. The American Robin (Turdus migratorius), for example, leaves Alaska and most of Canada for the winter and ranges as far south as Guatemala. Some years, more birds retreat, and some may go farther south in other years. Is this a stepwise migration, with all birds moving southward to some degree, or are some robins traveling various distances and others not moving at all? The term partial migration covers many variations in migratory behavior.

SHORT-DISTANCE MIGRATION The entire population moves out of the breeding range, but usually the distance between the limits of the breeding and winter ranges is less than 2000 km. McCown’s Longspur (Calcarius mccownii) and Chestnut-collared Longspur (C. ornatus) have overlapping breeding ranges in the northwestern Great Plains and southern Canadian Prairie Provinces, and both winter from Oklahoma and Texas south into northern Mexico. They have to move south in winter to reach latitudes where the grasslands and deserts are not covered with snow. Similarly, the Rusty Blackbird (Euphagus carolinus), which breeds in bogs and muskeg swamps from maritime Canada to Alaska, vacates that vast range for the central and southern United States from the Atlantic to about the 95th meridian, where swamps, wet woodlands, and pond edges are less likely to be frozen (Avery 1995).

Recent tracking of Arctic Terns has revealed that, for some, the extent of winter travel is even greater than previously believed. Some terns from Greenland spend part of the austral summer far in the Indian Ocean. (redrawn with permission from Egevang et al. 2010)

LONG-DISTANCE MIGRATION The entire population moves, usually more than 2000 km, often to a different continent. As with the short-distance migrants, their destinations are determined by the reliable availability of food to sustain them through winter. The most extreme example is the Arctic Tern (Sterna paradisaea), which nests around the globe almost entirely above latitude 59° N (some almost at 84° N) and winters at sea south of 58° S, primarily at the edge of the pack ice around Antarctica; some birds from Greenland colonies go south through the Atlantic and spend part of the winter in the Indian Ocean before returning to the South Atlantic. They travel 80,000 km each year and spend several months at both ends of their migration in perpetual daylight (Egevang et al. 2010). Unlike most terns, Arctic Terns feed substantially on crustaceans as well as fish; in their high-latitude nesting and wintering areas crustaceans are abundant, and the Arctic Tern avoids competition with several close relatives in the lower latitudes of all oceans (Hatch 2002).

ALTITUDINAL MIGRATION Some birds, as already noted, make an altitudinal migration. Appalachian and some other mountain populations of the Dark-eyed Junco (Junco hyemalis) descend to lower elevations in winter, while more-northerly juncos move south; where two junco subspecies occur together during winter in the Great Smoky Mountains, most local breeders settle higher on the slopes than the birds from farther north (Rabenold and Rabenold 1985). In Bolivia, where from the Andes to the Amazon lowlands 34 flycatchers are found year round, 15 shift their elevational range 500 m or more, one as much as 2000 m. Other flycatchers there are partial migrants, in which a portion of the population shifts downward for winter, either withdrawing from the upper elevations of the breeding range or moving to elevations below the breeding range (Chesser 1997).

In addition to the distance traveled by each species, the annual movements must be considered in terms of which population goes where. For any migratory bird, competition for winter resources is reduced if the population is spread through the largest possible winter range. This can be achieved in different ways. Among Northern Hemisphere birds, all may move southward stepwise, more or less the same distance, rather than to the same destination, so that the more northerly breeding population remains farther north during winter even after all birds have shifted southward. Other birds, such as the Common Yellowthroat (Geothlypis trichas), have a leapfrog migration, in which the northernmost population goes farthest south; migrant yellowthroats, in fact, also leap over a resident population in California and the southeastern states. Birds that have a wide east-west range may move due south—if geography allows—in a parallel pattern, or they may cross each other.

In many species, birds from different parts of the breeding range mix during winter. This is especially likely to take place where the winter range is much smaller than the breeding range, as for the warblers that nest in the vast boreal forest of North America and winter in Central America and islands of the Caribbean.

In other species, however, there is a strong link between the breeding and winter ranges of certain populations, called migratory connectivity, discussed at length in chapter 3. Long-billed Curlews (Numenius americanus) nesting in Oregon, Nevada, and Montana winter almost entirely in separate regions. Oregon birds go to the Central Valley of California; a few Nevada birds go there as well, but more go to Baja California; and Montana birds winter on the Mexican Plateau, in the Texas Panhandle, and near the Laguna Madre on the Gulf of Mexico (Page et al. 2014).

While selection pressures may be significant in any part of the annual cycle, for most birds the survival challenges of winter are greater than in the breeding season, which is usually much shorter. Among long-distance migrants, winter selection has the greatest evolutionary impact on species with high migratory connectivity, when the breeding population of a species in one region, already somewhat genetically isolated, also winters together, thereby subjecting all birds to that season’s local pressures as well. Breeding populations that disperse in winter and meet conspecifics from other regions will be less affected by distinctive selection pressures, because not all members of a regional breeding population will be exposed to the same selection factors during any winter.

Impacts of winter selection may be especially strong on birds that spend more of the year at their wintering sites than in their breeding range, such as Arctic-nesting shorebirds and geese. Even many birds breeding in temperate latitudes spend most of the year where they winter, although winter, as we think of it with colder weather, may be at most only a fraction of their time there. Long-billed Curlews that nest in Oregon return to the Central Valley of California as soon as mid-June, remaining there until the following March or April (Page et al. 2014). The same is true for many birds going to tropical regions where there is no cold at all: Kentucky Warblers (Geothlypis formosa) and Louisiana Waterthrushes (Parkesia motacilla) that nest in the southeastern United States arrive in Veracruz, Mexico, by late July or early August and stay until late March (Rappole 1995:79).

Kentucky Warblers winter in Veracruz, Mexico, from late July until late March, making this their home for far more of the year than their breeding range in the United States.

Body size is one of the most widespread aspects of winter selection pressure. Within any species of bird (and many other animals), overall body size varies among populations based on latitude, usually with those of colder regions being larger, because larger bodies retain heat more efficiently. Nonmigratory Hairy Woodpeckers (Dryobates villosus) resident in Alaska are half again larger than those in Mexico. In migratory Canada Geese (Branta canadensis), however, the smallest races nest farthest north. They have a leapfrog migration, with the most northerly races wintering farthest south, on the Gulf Coast and in Mexico. For them, there is evidently a selective advantage to being smaller, so their body dissipates heat more easily during the many months of the year they spend in relatively warm latitudes.

OTHER SEASONAL MOVEMENTS

Not all birds that leave their breeding area in winter have a regular latitudinal or altitudinal migration. In some species, notably certain seabirds, populations disperse broadly from their colonies over a wide area of water or coast. Adult males usually remain closest to their nesting site and are the first to return at the end of winter, while adult females and young birds move farther; young birds sometimes eventually settle at other colonies. In passerines, winter dispersal may be only a few tens of kilometers within the breeding range as birds search for better short-term feeding sites in places that may not be suitable for breeding (Newton 2008:513–514).

Dispersal may be an appropriate response to some local winter conditions but not others. Among Herring Gulls (Larus argentatus), young birds from around the Great Lakes disperse during their first winter throughout eastern North America from James Bay to the Gulf Coast and Mexico, while first-year birds born on the Atlantic coast move south as far as the Gulf, and the birds of the St. Lawrence region are resident (Eaton 1934).

Some northerly birds can survive most winters where they breed but must move if their food source fails. Irruptions bringing these birds sometimes far south of their usual range happen every few years, for some species with more regularity than for others. Archetypal irruptive species include finches like crossbills (Loxia spp.) and redpolls (Acanthis spp.), and the Snowy Owl (Bubo scandiaca). Their cyclical foods precipitate these movements: tree seeds for the finches, and rodents, especially lemmings, for the owl.

Trees of the boreal forest produce different quantities of seeds each year, usually on a two-year cycle. A study correlating 30 years of Christmas Bird Count tallies, breeding bird surveys, and cone and seed crop data in North America found that the irruptions of most finches occur in years when poor crops follow a year with a large conifer seed crop. For Common Redpolls (Acanthis flammeus) and Black-capped Chickadees (Poecile atricapillus) in the West, however, irruptions followed directly after large seed crops, not after subsequent crop failures. In no cases were irruptions linked with milder winters that led to higher overwinter survivorship, nor did severe winters result in abandonment of normal winter range, or warm conditions in the breeding season generate greater reproductive success (Koenig and Knops 2001). When seeds are easily available, bird populations usually increase and remain; when the food source drops off or has been consumed, these expanded bird populations must move elsewhere.

Winter finches—the various species that in some years appear in numbers well south of their breeding range—shift their diet in late summer or autumn, when seed crops are maturing; their movements may begin long before winter if local resources are poor (Bock and Lepthien 1976). Crossbills, which feed on the seeds in pine and spruce cones, will wander widely when a cone crop diminishes or fails. As flocks search for food over long distances, they may even nest in midwinter if they find a reliable source. Both species of redpolls depend on birch seeds and may also move far from one breeding area to another. Common Redpolls banded one winter in New England appeared in Alaska the next winter, and one banded in Michigan was recovered a year later near Okhotsk in eastern Siberia, 10,200 km distant (Troy 1983). Another found in Belgium one winter was in China the next, 8350 km away (Newton 2008:541). Where any of these birds nested during the intervening summer is unknown.

Other small birds of the northern forest are also irruptive, often in synchrony, even though their diets do not overlap. General environmental conditions in these forests may lead to good or poor years for both plants and insects. An analysis of Christmas count data from 1959 through 1997 showed strong two-year synchrony, especially in eastern North America, among Red-breasted Nuthatch (Sitta canadensis), Black-capped Chickadee, and Pine Siskin (Spinus pinus)—but only during some decades. Invasions of some other irruptive species, however, from the insectivorous Golden-crowned Kinglet (Regulus satrapa) to the frugivorous Bohemian Waxwing (Bombycilla garrulus) and the bud-and seed-eating Purple Finch (Haemorhous purpureus), did not correlate strongly with the movements of any other species (Koenig 2001). Another study found little correlation between North American irruptions of several highly synchronized species in boreal forests and populations of the same species in similar forests at high elevations in the western United States (Bock and Lepthien 1976). It will take even longer-term studies, including more data on changes in food sources in different parts of the breeding range, to reveal the causes and patterns of these irruptions.

Snowy Owls preying on Arctic rodents—which themselves respond to food availability on the ground—go through cycles of abundance and die-off averaging three to five years. A study of Snowy Owl nesting productivity from 1994 to 2011 on Bylot Island in the Canadian High Arctic and Daring Lake, Northwest Territories, in the Low Arctic found that winter irruptions were correlated with the most successful breeding seasons, which create the greatest potential competition for food the following winter. Since young Snowy Owls do not themselves breed for several years, they have less incentive to remain near their breeding range. By going south they spend at least their first winter in milder conditions, perhaps with more easily available prey, and with less competition from adults (Robillard et al. 2016).

The Snowy Owl is the most conspicuous and regular of owls in cyclical irruptions, but it is not unique among northern rodent predators. When necessary, Boreal Owls (Aegolius funereus) and Northern Hawk Owls (Surnia ulula) move in winter, mainly within the boreal forest zone. Short-eared Owls (Asio flammeus) and Long-eared Owls (A. otus) wander south in irregular numbers each winter, returning north the following spring and settling wherever voles are plentiful. Larger raptors that prey on snowshoe hares (Lepus americanus) move in response to the hare’s roughly 10-year cycle; Great Horned Owls (Bubo virginianus) and Northern Goshawks (Accipiter gentilis) may irrupt for as many as three winters running while hare populations are low (Newton 2008:579).

MIDWINTER AND FACULTATIVE MIGRATION

At middle and high latitudes, many short-distance migrants live where weather may change suddenly and dramatically, with freezing temperatures, snow, and ice transforming the landscape from one day to the next and eliminating food or preventing birds from finding it. Facultative migration, the decision a bird makes to move weeks after it has finished its normal autumn migration and reached a destination within the species’ usual winter range, may come irregularly rather than annually, in response to unpredictable conditions during the severest part of an exceptional winter. Some individual birds capable of facultative migration may not experience the need for it during their lifetime.

These sudden transformations of the landscape may provoke substantial flights of birds that are otherwise scattered across a broad region. On December 28, 1964, 20,000 Eurasian Skylarks (Alauda arvensis) were counted passing westward over the Axe Estuary in Devon, England, and on January 30, 1972, a group of 10,200 Northern Lapwings (Vanellus vanellus) flew over Portsmouth, Hampshire, in 2.5 hours. There are no comparable figures from North America. In Europe, this mass evacuation phenomenon has become less frequent in recent decades as winters have grown milder (Newton 2008:469–470). But anywhere that water bodies may freeze rapidly in midwinter, ducks or geese first become concentrated in the diminishing area remaining open, and then they disappear overnight—with an influx farther south where water is open. Often, also flying at night, these birds will return a few days later when conditions become more favorable, thereby saving themselves from continued competition in the densely packed area where they found refuge with other waterfowl already present.

The Eurasian Golden-Plover (Pluvialis apricaria) is normally a short-distance migrant, with birds from Scandinavia wintering in northwestern Europe, where they feed primarily in pastures and agricultural fields, foraging for earthworms and insects. Plovers from Swedish Lapland tracked in 2011–2012 with light-level geolocators began their winter in the Netherlands, Belgium, and northwestern France. When temperatures in some parts of that region dropped below freezing between January 30 and February 1, the plovers there moved within a day to southern Spain and Portugal, and some then moved in the next day or two to Morocco. They remained at these locations until March, while other tracked plovers that were in England and western France, where the weather was milder, stayed there all winter (Machin et al. 2015).

Facultative migration has been well studied in a few passerines. In the area around Phoenix, Arizona, Yellow-rumped Warblers (Setophaga coronata) commonly winter in isolated riparian valleys with permanent water that are surrounded by the Sonoran Desert and agricultural lands. Here, they feed on insects on foliage, but occasional cold fronts that generate freezing temperatures, high winds, and hail may cause the leaves to fall from all trees. When this happens, there is a substantial decrease in warbler numbers around Phoenix—at one site where the warblers were counted all winter, from 285 birds/40 ha to 34 birds/40 ha in 11 days—with a corresponding influx in northern Sonora, Mexico, 325 km southeast, which had 11 birds/40 ha and 10 days later had 90 birds/40 ha (Terrill and Ohmart 1984).

Further evidence of facultative migration in Yellow-rumped Warblers, as well as proof that the birds migrate at night, as they do in their regular migration, comes from Florida, where midwinter counts of warblers killed at night by hitting a radio tower in Leon County correlated with cold fronts. During seven mild winters, the number of Yellow-rumped Warblers killed between December 15 and February 20 was as low as 5; in the severe winter of 1962–1963 it was 264 (Terrill and Crawford 1988). In that winter, this species was common in Jamaican lowland forests and mangroves into April, while in other winters it is conspicuously absent (Gochfeld 1985).

Experiments with two regular facultative migrants showed how this type of migration is triggered. Dark-eyed Juncos held in captivity during winter showed little or no restlessness at night when given large amounts of food. But when the quantity was restricted, their nocturnal activity paralleled migratory restlessness typical of autumn. This could be induced during December and January, but after late January, the migratory behavior could not be reactivated with food reductions comparable to what birds had received earlier in the winter. Similarly, Garden Warblers (Sylvia borin) deprived of food in midwinter will show the same nocturnal activity as during their autumn migration season, and they will cease this activity as soon as food is returned and they begin to regain weight. In winter, however, unlike in autumn, they will not seek to migrate again after they have returned to normal weight (Terrill 1990).

For some birds, facultative migration may continue until the end of winter. Altitudinal migrants, which are much closer to their breeding sites in winter than are latitudinal migrants, may go back and forth depending on weather conditions. Yellow-eyed Juncos (Junco phaeonotus) in the Chiricahua Mountains of southern Arizona nest as high as 2560 m and commonly descend to winter 8 or 10 km away where the elevation is 1000 m less. In late winter, they return to higher elevations to stake out or reclaim territories even though they will not begin nesting for another several weeks. After storms, however, some individuals depart again for their wintering area at lower elevations as soon as the next morning. Within a few days, they are back in their breeding territory (Horvath and Sullivan 1988).

MAJOR MIGRATION SYSTEMS

On every continent, substantial numbers of species migrate to avoid winter at high latitudes. Some 318 North American landbird species move to the Neotropics, from the Palearctic 185 go to Africa, and South Asia receives 338 species from the eastern Palearctic. To these numbers must be added the many migratory species that stay in temperate latitudes through the winter. In South America, at least 220 species of all kinds move northward for the austral winter, while in Australia about 40% of landbirds migrate, and the movements of other birds are governed by irregular patterns of rainfall, resulting in more nomadism than classic migration. In Africa’s midbelt south of the Sahara, seasonal rains rather than significant shifts in temperature also govern most migration; 40% of West African landbird species, for example, may move within the tropics (Rappole 2013:260–269). Farther south in Africa, many birds move north during the austral winter. In each region, the migratory response of landbirds to winter reveals a different pattern set by climate, geography, and evolutionary history. Similarly, seabirds that touch land only to breed move the rest of the year through different parts of the global ocean to reach areas where seasonal resources are abundant; this is most striking in the many shearwaters and petrels that breed at high latitudes in the Southern Hemisphere and avoid winter there by migrating to the North Atlantic and North Pacific, where it is summer.

NEARCTIC/NEOTROPICAL

Among landbirds of North America, most of the migrants to the Neotropics breed in forests. When foliage that sustains their arthropod prey falls off the trees in deciduous forests and cold reduces similar prey in coniferous forests, these birds migrate to tropical and semitropical latitudes where arthropods remain reliably available—even as some of these birds broaden their diet to include fruit and nectar, which are scarce or absent farther north. Most of these forest birds concentrate in the latitudes where they may have originated: between southern Mexico and northern South America, as well as on Caribbean islands, seeking habitat that is at least superficially similar to the woody vegetation they use