Animals and Ecological Science

Oxford Handbooks Online

Animals and Ecological Science Anita Guerrini The Oxford Handbook of Animal Studies (Forthcoming) Edited by Linda Kalof

Subject: Political Science, Political Theory, Comparative Politics Online Publication Date: Feb DOI: 10.1093/oxfordhb/9780199927142.013.25 2015

Abstract and Keywords

Ecological science, which studies the relationships between organisms and their environments, developed from natural history. Aristotle’s teleological chain of being and detailed description modeled natural history until the eighteenth century. Linnaeus and Buffon replaced Aristotelian categories with new criteria for classification, leading the way to Darwin’s evolutionary theory. Darwinian evolution depended on environmental factors and led to the birth of ecological science by the end of the nineteenth century. The ecosystem concept emphasizes populations and systems rather than individuals. Case studies, of wolves and fish show the range of modern ecological science. Anthropogenic changes to the environment have led to extinction and endangered species. Attempts to meliorate human influence include rewilding and synthetic biology.

Keywords: ecological science, chain of being, ecosystem, endangered species, evolution, extinction, natural history, rewilding, synthetic biology, taxonomy

Introduction

The chapter examines animals as objects of scientific study in the ecological sciences under three broad headings. Before there was ecological science, there was natural history, while today some believe that the future of ecological science is in what has been called de-extinction or synthetic biology. Between these two extremes, I look at some current practices in ecological science, with case studies of wolves and fish. The following section of the volume looks at animals in ecosystems.

From Natural History to Ecology

Although Aristotle (384–322 BCE) was not the first to regard animals as subjects of inquiry rather than as commodities, he was the first Western philosopher to do this systematically. His works on animals, particularly History of Animals, Parts of Animals, and Generation of Animals, established a science of natural history that endured until Darwin and in some ways persists today. Historia (Greek ἱστορία) originally meant simply an “inquiry” or an “investigation,” or an account of such an inquiry. It did not imply the passage of time. Aristotle’s History of Animals offered detailed descriptions of all animals known to him. Unlike his mentor Plato, Aristotle was no armchair philosopher, and he took every opportunity to observe every animal he could: wild and domestic, native and exotic, terrestrial and aquatic. He investigated morphology, habitat, behavior, and what he called “manner of life”; what parts were the same and what were different; how they ate and reproduced. He noted natural kinds and attempted various classifications. Broad groupings seemed obvious: birds were different from fish. Some animals had two feet, some four, others none. Some animals were “blooded”; some, like insects, were not. But when Aristotle began to look at generation, he found categories that cut across others and that seemed to fit a hierarchical system based on degrees of perfection as measured by degrees of natural heat. Thus warm-

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blooded viviparous animals were “hotter” and “more perfect” than oviparous animals, and so forth, down to those animals that he believed produced larva rather than eggs. This hierarchical system, later known as the “chain of being” or “ladder of nature,” proved to have remarkable staying power in Western thought. The chain of being was not only hierarchical but full, including every animal (and plant) that could be created. It was also unchanging, so that species were fixed in time and space. And it was teleological: nature always worked toward a purpose.1

From the outset, the natural history of animals did not consist merely of passive observation. Aristotle dissected many dead animals and a few living ones. He collected and preserved specimens. These remained essential practices for the science of natural history, as did recording observations in words and pictures. Natural history overlapped with other uses of animals: collections of exotic animals in menageries conferred prestige on their owners but also provided opportunities for naturalists to observe new species, and hunters and fishermen often provided materials for study. The Roman physician Galen (c. 129–210 CE) used animals in anatomical studies to learn about human function, but at the same time, he also learned about animals. Naturalists from antiquity to the nineteenth century followed Aristotle’s example and collected, dissected, and observed.

Christians, Muslims, and Jews adopted Aristotle’s hierarchical concept of nature and scientists still refer to “higher” and “lower” animals. Beginning with the influx of New World animals to Europe in the sixteenth century, however, the chain of being began to fall apart. For example, the Swiss naturalist Conrad Gessner (1516–1565) did not quite know what to do with the armadillo, and he strained to fit it into a known niche on the chain of being. As translated by Edward Topsell (1572–1625) a half century later, the “Tatus or Guinean Beast” (“Guinean” in this era simply meant “foreign”),

is brought for the most part out of the new-found world, and out of Guinia, and may therefore be safely conveyed into these parts, because it is naturally covered with a harde shell, devided and interlined like the fins of fishes, outwardly seeming buckled to the backe like coat-armor, within which, the beast draweth up his body, as a Hedghog doth within his prickled skin; and therefore I take it to be a Brazilian Hedghog.2

Gessner’s Historiae animalium (1551–1558) was one of a number of encyclopedic natural histories of the Renaissance that drew on the ancient Roman Pliny (23–79 CE). Pliny’s very popular Natural History surveyed all that was known about nature, mingling direct observation with a variety of textual sources of varying credibility. Aelian (170–230 CE) followed this model, as did Christian writers such as Isidore of Seville (560–636), who included a section on animals in his Etymologia, and Albertus Magnus (c. 1200–1280), whose De animalibus both summarized Aristotle’s animal works (which Albertus reintroduced to the West) and displayed his own observations, particularly on the falcon.

Gessner and his contemporaries, such as Ulisse Aldrovandi (1522–1605), made no attempt to classify beyond very general categories. The lack of consensus about classification among naturalists is evident in cabinets of curiosities, assembled in this period as physical counterparts to Renaissance natural history texts. Cabinets served as prototypes for natural history museums, which emerged at the end of the eighteenth century. A cabinet belonged to an individual and reflected that person’s tastes and interests. Usually a single large room, it functioned as a naturalist’s workplace and as a site of display, open or not to spectators. Collectors mingled natural history and antiquities, natural objects and made objects, using surprising juxtapositions to produce particular effects: aesthetic, moral, or philosophical. Unusual specimens and natural anomalies were particularly prized, but cabinets also documented the ordinary course of nature. Illustrated catalogs mapped the collections. For example, the cabinet of Italian apothecary Ferrante Imperato (1550–1625) featured many preserved animal specimens, with an emphasis on the rare and unusual—the diarist John Evelyn reported seeing chameleons and “an extraordinary greate Crocodile.”3 Preservation methods included drying and “wet” preparations in jars with some kind of preserving fluid, as well as taxidermy. Late seventeenth-century works of natural history, such as Mémoires pour servir à l’histoire naturelle des animaux (1671–1676) of the Paris Academy of Sciences served as a kind of paper cabinet or paper menagerie, since it consisted of animals from Louis XIV’s menageries.4

The idiosyncratic organization of cabinets and such works as the Mémoires reflected continued debate about the proper criteria for classification. The ideal, a system that displayed the order of nature, seemed increasingly out of reach. Aristotle had attempted and failed to establish such a natural system. The naturalist John Ray (1627–1705), who edited the comprehensive natural history of birds of Francis Willughby (1635–1672), attempted at the end of the seventeenth century to outline a natural classification of animals, but it was generally viewed as too complex to

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be useful. The discovery of the sexuality of plants, which emerged from the work of a number of late seventeenth- century naturalists, provided a key to the classification of both plants and animals. Swedish botanist Carl von Linné (Carolus Linnaeus, 1707–1778), classified plants according to sexual parts in his Systema naturae (1735), which also presented a scheme for classifying animals, organizing them in six broad classes: quadrupeds, birds, amphibians, fish, insects, and worms. The tenth edition in 1758 established the now standard binomial nomenclature of genus and species. Linnaeus aimed to establish order rather than to reproduce nature’s plan, but he saw that order as a revelation of God’s design.5

Georges-Louis Leclerc, Comte de Buffon (1707–1788), compiled the most comprehensive eighteenth-century work of natural history in the 44-volume Histoire naturelle (1749–1788). Buffon viewed himself as a new Pliny who would write a natural history of everything, and as supervisor of the royal botanical garden in Paris and its menagerie, he presided over a worldwide trade in plants and animals.6 With his collaborator Louis-Jean-Marie Daubenton (1716– 1800), volumes 3–15 on quadrupeds established comparative anatomy as an essential component of natural history. Buffon also took advantage of technical developments in engraving and printmaking over the previous two centuries in his detailed illustrations.

Buffon acknowledged in his preface that we cannot pretend to be able to understand all of nature’s complexity and abundance. He dismissed much of the work of earlier naturalists and claimed a method of observation and comparison of numerous individuals derived from the English philosopher Francis Bacon (1561–1626), although he also referred to both ancient and more recent works such as the Paris Academy’s Mémoires. Perhaps most importantly, he disregarded any religious framework; although nature revealed a design, it did not come from God. Buffon’s first discovery was, he noted, “perhaps humiliating to humans: it is that we must classify ourselves as animals.”7 He went on to condemn all current classification schemes as being artificial and incomplete, displaying “an error of metaphysics,” what we might call a category mistake. The systems for classifying animals were, he said, even worse than those for plants.8

Buffon sought a natural system based on close observation of all characteristics of an organism. Such a system would not be as complete and complex as that of Linnaeus, but it would, he believed, be truer to nature. In the volumes on quadrupeds (what we would call mammals, following Linnaeus), Buffon and Daubenton organized them first into domestic and wild, then into local and exotic, with further divisions according to teeth and other criteria. But over the 40 years of the Histoire naturelle, Buffon added the critical concept of time to natural history. Acknowledging studies of fossils that had begun with Nicholas Steno (1638–1676) in the previous century, Buffon came to emphasize contingency and historical process in nature, arguing that present life forms can be explained by their history. In addition, Buffon acknowledged what Steno and others had been reluctant to recognize: the fact of extinction. Not all life forms survived; nature’s plan, whatever it was, included imperfection and dead ends. In Epoques de la nature (1778), Buffon presented a story of natural development over time that greatly extended the traditional Christian time frame based on biblical chronology. Together with Linnaean classification, Buffon laid the groundwork for the development of a secular science of natural history in the nineteenth century. Buffon’s King’s Garden, which the French Revolution re-created as the Paris Museum of Natural History, became a central site for the development of this new science.9

Museum collections of animals in the nineteenth century, based on voyaging and collecting over the previous 200 years, engendered the professionalized and specialized natural history of animals that in turn gave birth to a number of new sciences, including ecology. Historians have traced the development of evolutionary theory from Georges Cuvier’s studies of fossils at the Paris Museum of Natural History in the 1790s to Darwin’s 1859 Origin of Species, a development that was neither smooth nor inevitable. The work of numerous naturalists, artists, collectors, curators, anatomists, and experimenters contributed to Darwin’s theory and its elaboration in the second half of the nineteenth century. Many of the same individuals contributed to the beginnings of the science of ecology, which historian Lynn Nyhart attributes to “taxidermists, zookeepers, school teachers, museum reformers, amateur enthusiasts, and nature protectionists.”10

The science of ecology, defined by Ernst Haeckel (1834–1919) in 1866 as the relationship of organisms with their environment, is a direct consequence of Darwin’s ideas of animals’ adaptation to differing environments. Unlike early modern naturalists who focused on individuals—since often they only had knowledge of a single specimen of a particular animal—Darwin’s evolutionary theory operated at the level of populations. This disappearance of the individual animal would have important implications for the ecological study of animals.

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Animals and Ecological Science: Two Case Studies

The modern study of animals that may be included under the umbrella of “ecological science” spans a gamut from broad and highly mathematical population studies to intimate studies of animal behavior. A popular notion of ecological research pictures rugged scientists observing nature in the wild, from a distance. There is some truth to this notion, but ecological research can also be as invasive and manipulative as other types of animal research. Much ecological work is now done in laboratories and with computer models, but it continues to be distinct from other biological sciences in its persistent engagement with field work. Early ecologists tried to capture this distinctiveness in varied titles; around 1900, Frederic Clements called himself an “outdoor physiologist”; in the 1930s Victor Shelford referred to ecology as “scientific natural history.”11

The concept of the ecosystem was first enunciated by British ecologist Arthur Tansley in 1935, but the idea had been around for a while. Whether nature was an organism, as Clements argued; a community, as Charles Elton maintained in his influential book Animal Ecology (1927); or a complex biogeochemical system of feedbacks and nutrient cycling, as G. Evelyn Hutchinson and later Eugene Odum demonstrated, it became increasingly clear that ecological science was one of systems rather than of individuals.12

Wolves

Although Romulus and Remus were said to have suckled on a she-wolf, wolves have more often been feared as predators of humans and symbols of an unknown and uncivilized world. In the Middle Ages, the taming of the wolf of Gubbio by St. Francis reflected this widespread fear of wolves in particular and the wilderness in general. In the eighteenth century, the bête féroce of the Gévaudan—finally revealed to be a wolf—terrorized a region in the Massif Central of France for a year.13 Wolves had become extinct in much of Western Europe by the end of the nineteenth century. In the early years of westward expansion in the United States, wolves were obstacles to be extirpated, but by the late twentieth century they had become symbols of lost wilderness and human hubris. In Eurasia and the United States, modern wolves have a mixed status as both an exotic animal in a zoo and a wild and not necessarily welcome native animal.

In the United States, grey wolves were among the first animals to be listed after passage of the Endangered Species Act in 1973 (the Bern Convention of 1979 performs a similar function in the European Union). Long considered to be vermin, wolves in the lower 48 states were confined to a small area in northern Minnesota and Michigan’s Upper Peninsula by the 1930s. Although the noted wildlife ecologist Aldo Leopold (1886–1948) had noted the “fierce green fire” die out in the eyes of a wolf he shot in Arizona in 1909 (as he recounted in his 1944 essay, “Thinking Like a Mountain”), he continued to hunt wolves and to advocate their slaughter for the next 20 years.14 Only in the early 1930s, when wolves had been nearly hunted, trapped, and poisoned out of existence, did he begin to recognize the role of predators in ecosystem maintenance. His 1933 book Game Management, for many years the main textbook on the subject, revealed in its title two enduring characteristics of human relationships with, and study of, wild animals: they were “game” for the use and sport of humans, and they required human management.15 The management context remains prominent in research on wild animals. Unlike laboratory animal research, ecological research is often applied research, particularly in the United States, as reflected in funding sources that include the Bureau of Land Management, the Fish and Wildlife Service (at the federal and state levels), and the Department of Agriculture (which oversees the Forest Service).16

By the 1980s, wolves were on the rebound in the United States and, owing to similar conservation measures, in parts of Europe as well. Wolves trickled across the border into the United States from Canada, and in a controversial measure, Canadian wolves were reintroduced into Yellowstone National Park and areas of central Idaho in 1994–1995. The reintroduction of wolves to areas that had not seen them in over half a century continues to raise many issues, both scientific and political, as well as a significant public response. In ecological terms, wolves are known as keystone predators, whose influence on an ecosystem extends far beyond their immediate impact on specific prey. The presence of such keystone species triggers “trophic cascades” of nutrient circulation, affecting a number of species throughout an ecosystem.17 For example, Ripple and Beschta recently demonstrated that the reintroduction of wolves to Yellowstone diminished elk populations, which in turn allowed aspen and cottonwood trees, beavers, and a number of other species to increase in numbers.18

Idaho wolves soon crossed the border into the states of Oregon and Washington, and in December 2011 an

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Oregon wolf known as OR-7 crossed the southern border of Oregon into California. The last known wolf in California had been shot in 1924. Scientists from the Oregon Department of Fish and Wildlife have tracked OR-7’s travels with a GPS collar he has worn since early 2011. Radio tracking, first used in the 1950s, has been used to acquire data that is then used on a metascale in population studies and modeling, where the individual wolf disappears into the statistical model.19 Ripple and Beschta’s much-cited work on trophic cascades in Yellowstone did not look directly at wolves at all, but measured the effects of their reintroduction on other plant and animal species.20 Although tracking allows wolves to be studied at a distance, wolves must be found and trapped in order to attach the collars, and while in captivity they can undergo further manipulation, including measurement and blood sampling.

OR-7’s travels reflect the uneasy balance between individuals and communities in ecological research on animals as well as the particular emotional and political space that wolves occupy. In a well-known 1980 article, the environmental philosopher J. Baird Callicott explored this tension in terms of the philosophical incommensurability between animal liberation and environmental ethics. One focused on individuals, the other on systems and species. The science of ecology, he argued, overturned the idea that nature was a “collection of subjects” in favor of seeing it as “a unified system of integrally related parts.”21 But in the case of the wolf, the individual has not quite been subsumed into the system.

Wolves are among the most intensely studied animals in North America. The case in similar in Europe: after being hunted almost to extinction in the 1930s, wolves began to recolonize France in 1992 by crossing over the border from the Italian Alps, and have been subject to intense scientific and political scrutiny. A few years later, Polish wolves crossed the border into Germany, and wolves have recently been spotted in Denmark, the Netherlands, and Belgium. With their small and scattered populations as well as their status as keystone predators and charismatic megafauna, it remains difficult for either researchers or the public to see wolves solely in terms of anonymous populations.22 OR-7 has become internationally famous, with his own Wikipedia entry; similarly, researchers know each individual wolf in the Isle Royale pack in northern Michigan, further discussed by Michael Nelson and John Vucetich in the next section of this volume.

Behavioral ecologist Marc Bekoff, among others, has examined the social lives of wolves. Beginning in the 1970s, Bekoff compared the play behavior of wolves, coyotes, and dogs. He found that infant coyotes were more aggressive than either wolves or dogs, and that play behaviors played important roles in communication and social relations within the pack.23 In other work, he showed that aggressive behavior among wolves did not necessarily lead to dominance in the pack, and that wolves were more social animals than coyotes, who were much more solitary.24 Other ethologists have further elaborated the social interactions within a wolf pack. David Mech showed that the size of the wolf pack, long thought to be related to the size and availability of prey, was in fact regulated by other factors as well, including kinship relations and social interactions among the wolves.25 Mech and his colleagues examined data from a number of wolf studies in the United States and Canada over a period of more than 40 years.

The focused and long-term ecological study of wolves is echoed in work on other animals, mainly charismatic megafauna, such as elephants, lions, tigers, and great apes. Such attention can be seen as part of a long tradition of symbolic values as well as modern ecological concepts. Other species may play equally important roles in ecosystems but receive less public attention: fish are one example.

Fish

In contrast to the intense individual scrutiny of wolves, it is difficult for most people to see fish as individuals. Far from being charismatic megafauna, fish are nonetheless sentinels of climate change and ocean pollution, a critical food source, and strikingly diverse. They live in oceans and rivers, lakes and streams, in deep and shallow waters. Many species are game animals, and others have been domesticated to the extent of being intensively farmed. Ecological research on fish is correspondingly diverse, and takes place both in the wild and in laboratories.

“In the wild” can in turn be broadly interpreted. The Sierra Nevada Aquatic Research Laboratory near Mammoth Lakes, California is a natural setting that is heavily managed: Convict Creek is divided into channels, and a complex system of artificial streams allows for experimental manipulation of both native and nonnative fish. Research takes place in these streams, in the laboratories, and in the surrounding lakes. Since the mid-1800s, many lakes in the high Sierra have been stocked with trout for sport fishing, mainly with nonnative varieties, such

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as brook and brown trout. The California Department of Fish and Game aerially stocks lakes with fingerling trout each spring and summer; many of these lakes would naturally have no fish. This occurs in high-altitude lakes outside California as well. Researchers since the 1990s have documented the negative impact of fish stocking on native species of fish and on other animals, including the endangered mountain yellow-legged frog.26 This research includes studies not only of fish, but also of frogs, insects, fungus, birds, and phytoplankton, well illustrating the interconnectedness of this ecosystem. In perhaps the most dramatic demonstration, researchers eradicated all fish from a lake by means of gill-netting, a less toxic alternative to the usual fish management tool, the pesticide rotenone. The endangered frogs subsequently flourished.27

Nonvalued species, in this case nonnative trout, were therefore sacrificed (a standard term in animal experimentation) for a greater ecosystem good. Such species may be deliberately targeted or may be collateral damage from the collection of other species. Electrofishing is a common method of sampling to assess fish distribution and abundance in streams. The fish are stunned by an electric current, caught in a net, and then measured, sampled, or otherwise examined before being returned unharmed. However, recent studies have noted that electrofishing may not be entirely harmless to the fish, and that repeated electrofishing could cause physical and stress-related injuries to both targeted and nontargeted species.28 Some researchers have also questioned the utility of lethal sampling, particularly in the case of top oceanic predators, such as sharks, whose populations are already endangered. Hammerschlag and Sulikowski point out that such sampling would be out of the question for large terrestrial carnivores.29

Like wolves, some fish are tracked rather than caught. Tracking migratory fish such as salmon had long been sought to assess survival along the much-dammed Snake and Columbia Rivers in the US Pacific Northwest. The development of PIT (passive integrated transponder) tags in the early 1980s made this possible. Tiny electronic tags are injected either intramuscularly or into the body cavity of fish, which are then tracked via a series of antennas that pick up the tags’ electrical signal. Each individual fish is uniquely identified in a database. Millions of fish have been tagged since the 1980s, and their life cycles traced from river to ocean and back to the river.30

As these brief examples show, ecological research on animals ranges widely and encompasses both the field and the laboratory. Particularly in the case of charismatic megafauna such as wolves, research is deeply intertwined with social and political ideas of value and use. As indicators of the health of waterways and oceans, fish are increasingly studied. But mysterious outbreaks, such as sea star wasting syndrome, reveal how much remains to be learned about oceanic wildlife.31

Extinction Is Not Forever?

Early modern natural historians grasped the notion of extinction with difficulty. The idea that a specific animal could simply disappear violated a number of common beliefs. Plant extinction seemed even less plausible. Aristotle had argued that species were eternal. The great chain of being would surely collapse if any spaces occurred among its tightly packed rungs, and extinction implied that God, in creating the world, had somehow made a mistake. Yet, quite apart from the evidence of fossils, several animals had become extinct in historical times in Europe, particularly in Britain. Brown bears were extinct in Britain by 1000 CE and had retreated to remote areas of northern and eastern Europe by the seventeenth century. Wolves had become quite rare in Britain by 1500 and had disappeared altogether two centuries later. The European beaver had disappeared from England by 1550 and was found only in isolated pockets in France. The death of the last native European ox or aurochs in Poland in 1627 was widely noted at the time; the breed had been under the protection of the king of Poland for over a century.32

By the nineteenth century, the fact of extinction, if not its scientific or theological implications, had become widely accepted. The discovery at the end of the eighteenth century of the bones of mammoths and of the giant sloth that Cuvier named the megatherium provided convincing evidence of animals who no longer occupied the planet. These animals, unlike some fossils, had no living analogues. Extinction became a key concept for Darwin, who argued that species that could not adapt to changing environmental conditions would become extinct.

While Cuvier and Darwin established that extinction was a natural process, most modern extinctions are based on human activity, whether hunting or destruction of habitat. Several scientists and journalists have argued that we are presently at the beginning of a sixth mass extinction. Past mass extinctions caused the loss of half or more genera; during the last mass extinction at the end of the Cretaceous era, 66 million years ago, 75 percent of all 33

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species disappeared.33

The concept of endangered species—that is, species in danger of extinction—emerged in the mid-twentieth century. Building on work begun in the 1940s, the International Union for the Conservation of Nature and Natural Resources (IUCN), founded in 1948, established its Red List in 1964 to document rare and endangered species. Now on the web, it is constantly updated.34 The US Endangered Species Act focused particular concern on habitats, as did the Bern Convention. In addition, the CITES agreement (Convention on International Trade in Endangered Species of Wild Fauna and Flora), drawn up by IUCN and signed by 80 countries in 1973 (it now has 180 signatories), regulates the international wildlife trade with the aim of conserving endangered species of both plants and animals. It is administered by the United Nations Environment Program. All these programs aim to conserve existing species. But by the early 2000s, other ideas began to emerge surrounding the question of endangered animals and endangered habitats.

Rewilding

The idea of rewilding encompasses both the rehabilitation of landscapes and the reintroduction of particular species. While the reintroduction of extant species into places where they have become extinct is a long-standing practice—the reintroduction of wolves is one example—rewilding takes this another step.35 In 2005, ecologist Josh Donlan, then a graduate student, burst onto the scene as the lead author of a short report in Nature entitled “Re- wilding North America.” Coauthored with a plethora of heavy hitters including Dave Foreman, founder of Earth First, conservation biologist Michael Soulé, and evolutionary biologist Harry W. Greene, among others, “Re-wilding North America” made a radical proposal: to restore the lost megafauna of North America by bringing large wild vertebrates elsewhere to fill these lost ecological niches. In other words, mammoths, American lions, and cheetahs, and the ancient Camelops, all of whom disappeared at the end of the Pleistocene era some 13,000 years ago, could be replaced with analogous species including elephants, African lions and cheetahs, and camels.36

What one blogger called “bringing sexy animals back”37 created a sensation, with wide attention in the global press. In a more detailed account a year later, Donlan and his coauthors explained their reasoning: “Earth is now nowhere pristine”; therefore, re-creating missing ecological functions could be justified, even though it would introduce nonnative species. They suggested that the functional significance of megafauna had been undervalued and that therefore reintroducing them could help to arrest the “ecological chain reactions” that would lead to additional extinctions. One of the animals they cited in support of their argument was the gray wolf, noting that in its absence from the United States, species such as deer and elk had increased the predation of certain plants. In particular, the predation of young trees had led to declines in aspen and other desired trees and to the degradation of riparian areas, leading in turn to impacts on birds and other animals and plants. Citing the work of Ripple, Beschta, and others on trophic cascades at Yellowstone, Donlan and his coauthors argued,

The restoration of functionality from the reintroduction of wolves may even include a buffering of Yellowstone’s biodiversity to climate change…. Similarly complex but now extinct ecological roles for the dozens of lost Pleistocene predators and megaherbivores of North America would seem possible if not likely.38

Presenting this plan as “an optimistic agenda for twenty-first century conservation,” Donlan and his coauthors argued that “we can no longer accept a hands-off approach to wilderness preservation as realistic, defensible, or costfree.”39

While the “Pleistocene Park” Donlan envisages has not yet come to pass in North America, rewilding efforts have taken hold in widely varying places. In Siberia, scientist Sergey Zimov has joined native Yakutian horses, moose, and reindeer in a reserve with wisent (European bison) and musk ox, and plans to reintroduce native antelope and Bactrian camels in lieu of the extinct native camel. These herbivores appear to have had some success in recreating the grasslands that once dominated this area. He calls this reserve Pleistocene Park. The missing animal here is the woolly mammoth, who, in this climate, cannot be replaced by an elephant.40 Other parks are in various stages of development in Latvia, New Zealand, and Saudi Arabia.41 But what two ecologists recently called, with vast understatement, the “tricky issue” of species substitution is far from resolved.42

The best-known and most controversial of these parks is Oostvaardersplassen in the Netherlands. The brainchild of

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ecologist Frans Vera, the Oostvaardersplassen site dates back to 1968, when its land was reclaimed as a polder. Rather than being developed, it reverted to a wetland, and native birds began to appear. In 1982, the Dutch government decided to route a railway line around the site rather than through it, and the idea for Oostvaardersplassen began to take form. Vera aimed to recreate a Pleistocene landscape of grazing animals that could gradually also include forest; Vera’s much-debated theory about the Pleistocene landscape is that it consisted of mixed and shifting forest and pasture. In place of extinct wild Tarpans and aurochs, Vera substituted Konik horses from Poland (thought to be descended from Tarpans) and Heck cattle. German Heck cattle, developed by back-breeding in the 1920s to mimic the extinct aurochs, had been a favorite of Hermann Goering. Red deer, which are more like elk than deer, were added to the site in 1992. Since that time, a number of other wild animals, including graylag geese, white eagles and many other birds, roe deer, foxes and stoats, and a number of small rodent species have migrated to the site. Missing thus far are large predators such as wolves or bears. Wolves have recently been spotted in the Netherlands, and Vera is hopeful that they will make the journey to the reserve. Given the thousand-mile wanderings of OR-7, this is not improbable.43

The central problem of Oostvaardersplassen, as many commentators have noted, owes largely to this lack of predators. Because the animals at Oostvaardersplassen have been declared wild, they are not fed or otherwise managed, and only natural processes operate. Not all the animals survive the winters, and in particularly harsh winters, hundreds have starved to death. Because Oostvaardersplassen is a fenced reserve (of some 6000 hectares, almost 15,000 acres), the animals cannot seek other grazing grounds. The spectacle of starving animals has been quite controversial in the Netherlands, with questions asked in the Dutch parliament, two commissions of inquiry, and finally, an agreement that animals who looked to be unable to survive the winter would be shot, a death deemed more humane than starvation. The numbers of animals fluctuate widely; the numbers of Heck cattle have recently ranged from nearly 4000 in 2011 to under 3000 in 2013.44

If some, like New Yorker writer Elizabeth Kolbert, find Oostvaardersplassen “faintly ridiculous”; others, she admits, find it “inspiring,” and an active Rewilding Europe movement aims to “make Europe a wilder place,” with plans to rewild one million hectares by 2020 and “providing a viable business case for wild nature.”45 Like Donlan, the Rewilding Europe proponents view rewilding as a way to deal with rural depopulation as agriculture becomes more and more industrialized and more people move to cities.

De-extinction: The Jurassic Ark?

Michael Crichton’s 1990 science fiction novel Jurassic Park brought the idea of de-extinction to the attention of the public. The human genome project was just beginning, and the first successful mammalian clone, the sheep Dolly, was still several years in the future. Crichton, who prefaced his work with pages of realistic-looking genetic code, proposed that dinosaur genetic material found in insects preserved in amber could form the basis for re-creating dinosaurs and, presumably, other extinct animals. Scientists expressed skepticism that dinosaurs could thus be cloned: their DNA, even if preserved in amber, would be too degraded after millions of years to be viable.46 But, particularly after the successful cloning of Dolly in 1996, other more recently extinct animals began to get serious attention. In 2000, scientists in India proposed cloning the extinct Indian cheetah, the last specimen of which had been shot in 1953.47 At about the same time, Australian scientists began attempts to clone a thylacine or Tasmanian tiger (extinct since 1936), while scientists in Russia began the much more arduous task of cloning a woolly mammoth (extinct for over 10,000 years).48

Only in the past few years have the new genomic technologies known as synthetic biology become sufficiently developed that the prospect of de-extinction has become more than wishful thinking. Geneticist George Church, in his 2012 book Regenesis, cited the 2003 cloning of the recently extinct Pyrenean ibex or bucardo as evidence that “extinction [is] no longer forever.”49 Church is at the forefront of a group promoting the de-extinction of a number of species by genetic means. At the top of the list is the passenger pigeon, a species that was once abundant in the United States but became extinct in 1914. Critics point out that the cloned bucardo lived only for seven minutes, succumbing to lung malformations that had also, though less severely, affected Dolly.50

Enough genetic material remains in preserved specimens that the prospect of revival by genomic means is a possibility. Standard cloning techniques involve transferring the nucleus of a somatic cell of the extinct animal into an enucleated host egg cell. But the genetic material from preserved specimens of passenger pigeons and

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thylacines is incomplete and requires further manipulation and genetic engineering to become viable. DNA deteriorates after death, so the older the specimen, the more difficult the genetic reconstruction would be; cloning a woolly mammoth would be more difficult than the passenger pigeon, but Sergey Zimov is optimistic that it can be accomplished.51 Back-breeding is a lower-tech form of such engineering; 90 years ago, the Heck brothers back- bred cattle to resemble their idea of an aurochs. The Tauros project in Europe is a more sophisticated breeding program with the same goal.52

Geneticists also see possibilities in genetic engineering for improving currently endangered species. Theoretically, genetic manipulation could introduce disease resistance or even diversify small, inbred populations. In such contexts, rewilding takes on a new meaning, and ecosystem scientists, conservation biologists, ethicists, and legal scholars have debated the possible consequences of de-extinction and the genetic engineering of the environment. Animal welfare has not been at the forefront of this discussion. For example, little has been said about the 57 goats who were implanted with engineered bucardo eggs to yield one birth. Since cloning continues to be a risky and unpredictable procedure, even target species may have less than optimal outcomes.

Ecosystem scientists and conservation biologists point out that the environment is constantly changing, and that the niche that passenger pigeons, for example, occupied in the nineteenth century may no longer exist. The chestnut trees they favored are long gone, and other birds may now fulfill their ecological role. Habitat loss has been the major cause of modern extinction, and reintroducing formerly extinct animals will not in itself create habitat. Among many potential risks are the introductions or spread of diseases, unexpected species interactions, and invasiveness.53 Some are optimistic that these risks can be dealt with: Donlan points to successful eradications of invasive species on islands as an example of the increased human ability to manage wild populations. But he notes, “We are currently better at manipulating genomes than at rewilding landscapes.”54

The ethical implications, for animals, humans, and landscapes, are numerous. Some contend that de-extinction is a form of restorative justice, a way to undo the wrongs of the past. Others believe that de-extinction is mere hubris, and in the words of environmental activist George Monbiot, “lonely captivity is likely to be the fate” of animals produced by de-extinction.55 The legal and social implications are even more daunting. Could engineered organisms be patented? How will reintroduction be conducted and regulated? Who will fund such research (now it is largely funded through private foundations)? How will such reintroductions affect current protections of endangered and threatened species? Legal scholars Jacob Sherkow and Henry Greely point out that “current protection of endangered and threatened species owes much to the argument of irreversibility.”56 If extinction is not forever, will protections for existing animals be weakened or even disappear?

Even the most sober commentators acknowledge that the “gee-whiz” factor in de-extinction exerts a powerful pull. How cool would it be to see a mammoth? But venturing into such speculative realms has taken popular perceptions of animal ecology far from its roots in natural history and its concern for systems over individuals. The loss of species and the decline of biodiversity continue. The genetic tools of de-extinction may at some time in the future help to mitigate some of these losses, but animals still stubbornly resist their reduction to cells and genes, and ecological science still has much to learn about and from animals in the wild.

Conclusion

Aristotle would, I think, be intrigued and delighted to learn about modern ecology and its discoveries about animals. Convinced of the fecundity of nature, he would nonetheless find it difficult to abandon the philosophical principles of hierarchy and teleology that formed the basis of the great chain of being. Neither the idea of extinction nor the idea that extinct animals can be brought back would fit his system of values or his idea of science.

Modern ecological science has incorporated ideas from natural history into a comprehensive theory of the interactions of living nature. Although ecology has retained its connection with the field, the individual animal often disappears amid statistical models and more recently genetics. But connection with the field, along with a concern with applications, continues to distinguish the ecological sciences from other biological sciences. Entangled in policy and politics, the ecological study of animals will continue to be a practice embedded in broader ideas about the value and future of wild nature.

Notes: 1

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(1) The classic work on this topic is Arthur O. Lovejoy, The Great Chain of Being (Cambridge, MA: Harvard University Press, 1936). For a broad (and still unsurpassed) overview of premodern ideas of nature, see also Clarence Glacken, Traces on the Rhodian Shore (Berkeley: University of California Press, 1967).

(2) Edward Topsell, The Historie of Foure-Footed Beastes (London: William Jaggard, 1607), 705.

(3) John Evelyn, Diary, ed. E. S. de Beer, 2 (Oxford: Clarendon Press, 1955), 330–331.

(4) On these points, see Anita Guerrini, The Courtiers’ Anatomists (Chicago: University of Chicago Press, 2015).

(5) For further discussion of these points, see Paul Lawrence Farber, Finding Order in Nature (Baltimore, MD: Johns Hopkins University Press, 2000); Phillip Sloan, “Natural History, 1670-1802,” in Companion to the History of Modern Science, ed. R. C. Olby, G. N. Cantor, J. R. R. Christie, and M. J. S. Hodge (London: Routledge, 1990), 295– 313.

(6) E. C. Spary, Utopia’s Garden: French Natural History from Old Regime to Revolution (Chicago: University of Chicago Press, 2000).

(7) Georges-Louis Leclerc, Comte de Buffon, Histoire naturelle générale et particulière: avec la Description du Cabinet du Roy. Tome première (Paris: Imprimerie Royale, 1749), 12; my translation.

(8) Buffon, Histoire naturelle, 20.

(9) Farber, Finding Order in Nature, 20; Sloan, “Natural History, 1670-1802,” 304–306.

(10) Lynn Nyhart, Modern Nature: The Rise of the Biological Perspective in Germany (Chicago: University of Chicago Press, 2009).

(11) Keith R. Benson, “The Emergence of Ecology from Natural History,” Endeavour 24, no. 2 (2000): 59–62. The tension between field and lab is explored in Robert Kohler, Landscapes and Labscapes: Exploring the Lab-Field Border in Biology (Chicago: University of Chicago Press, 2002).

(12) Benson, “Emergence of Ecology.”

(13) Jay M. Smith, Monsters of the Gévaudan: The Making of a Beast (Cambridge, MA: Harvard University Press, 2011).

(14) Aldo Leopold, “Thinking Like a Mountain,” in A Sand County Almanac (New York: Oxford University Press, 1949). On Leopold’s thought, see Susan L. Flader, Thinking Like a Mountain: Aldo Leopold and the Evolution of an Ecological Attitude toward Deer, Wolves, and Forests, 2nd ed. (Madison: University of Wisconsin Press, 1994).

(15) Aldo Leopold, Game Management (1933; rpt. Madison: University of Wisconsin Press, 1987).

(16) On the history of the concept of biodiversity, see Robert Kohler, All Creatures: Naturalists, Collectors, and Biodiversity, 1850-1950 (Princeton, NJ: Princeton University Press, 2006).

(17) For a cogent explanation of trophic cascades, see Cristina Eisenberg, The Wolf’s Tooth (Washington, DC: Island Press, 2010).

(18) William J. Ripple and Robert L. Beschta, “Trophic Cascades in Yellowstone: The First 15 Years after Wolf Reintroduction,” Biological Conservation 145 (2012): 205–213.

(19) Tim Coulson, Daniel R. McNulty, Daniel R. Stahler, et al., “Modeling Effects of Environmental Change on Wolf Population Dynamics, Trait Evolution, and Life History,” Science 334 (2011): 1275–1278.

(20) Ripple and Beschta, “Trophic Cascades.”

(21) J. Baird Callicott, “Animal Liberation: A Triangular Affair,” Environmental Ethics 2 (1980): 311–338, at 321.

(22) Charismatic megafauna are defined as large animals (over 50 kg) who have widespread popular appeal.

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(23) Marc Bekoff, “Play Signals as Punctuation: The Structure of Social Play in Canids,” Behaviour 132 (1995): 419– 429.

(24) Marc Bekoff, “Mammalian Dispersal and the Ontogeny of Individual Behavioral Phenotypes,” American Naturalist 111 (1977): 715–732.

(25) See L. D. Mech and Luigi Boitani, eds. Wolves: Behavior, Ecology, and Conservation. (Chicago: University of Chicago Press, 2003), esp chap. 1, pp. 1–34; Paul A. Schmidt and L. David Mech, “Wolf Pack Size and Food Acquisition,” American Naturalist 150 (1997): 513–517.

(26) R. A. Knapp, “Non-Native Trout in Natural Lakes of the Sierra Nevada: An Analysis of Their Distribution and Impacts on Native Aquatic Biota,” in Sierra Nevada Ecosystem Project: Final Report to Congress, vol. III, Assessments, Commissioned Reports, and Background Information, 363–407 (Davis: University of California, Centers for Water and Wildland Resources, 1996).

(27) R. A. Knapp, P. S. Corn, and D. E. Schindler. “The Introduction of Nonnative Fish into Wilderness Lakes: Good Intentions, Conflicting Mandates, and Unintended Consequences.” Ecosystems 4 (2001): 275–278; R. A. Knapp, D. M. Boiano, and V. T. Vredenburg, “Removal of Nonnative Fish Results in Population Expansion of a Declining Amphibian (Mountain Yellow-Legged Frog, Rana muscosa),” Biological Conservation 135 (2007): 11–20.

(28) Frank M. Panek and Christine L. Densmore, “Electrofishing and the Effects of Depletion Sampling on Fish Health: A Review and Recommendations for Further Study,” in Bridging America and Russia with Shared Perspectives on Aquatic Animal Health, ed. R. C. Cipriano, A. W. Bruckner, and I. S. Shchelkunov (Landover, MD: Khaled bin Sultan Living Oceans Foundation, 2011), 299–308.

(29) Neil Hammerschlag and James Sulikowski, “Killing for Conservation: The Need for Alternatives to Lethal Sampling of Apex Predatory Sharks,” Endangered Species Research, 14 (2011): 135–140. See also Ben Minteer, James P. Collins, Karen E. Love, Robert Puschendorf, “Avoiding (Re)extinction,” Science 344 (2014): 260–261.

(30) “PIT Tag Information Systems (PTAGIS),” Pacific States Marine Fisheries Commission (PSMFC.org), n.d. http://www.psmfc.org/program/pit-tag-information-systems-ptagis accessed 17 November 2014.

(31) Dylan McDowell, “Environmental Drivers May be Adding to Loss of Sea Stars,” Breaking Waves 24 July 2014 http://blogs.oregonstate.edu/breakingwaves/2014/07/24/environmental-drivers-may-adding-loss-sea-stars/ accessed 17 November 2014.

(32) Wladyslaw Szafer, “The Ure-ox, Extinct in Europe since the Seventeenth Century: An Early Attempt at Conservation That Failed,” Biological Conservation 1 (1968): 45–47.

(33) See Elizabeth Kolbert, The Sixth Extinction (New York: Henry Holt, 2014).

(34) “The IUCN Red List of Endangered Species,” Red List. 2014.3 http://www.iucnredlist.org/ accessed 17 November 2014.

(35) Dolly Jørgensen, “Reintroduction and De-extinction,” Bioscience 63 (2013): 719–720.

(36) C. Josh Donlan, Harry W. Greene, Joel Berger, et al., “Re-wilding North America,” Nature 436 (2005): 913–914.

(37) Neil Chambers, “Josh Donlan on Bringing Sexy Animals Back via ‘Rewilding’,” treehugger.com, 24 December 2008, http://www.treehugger.com/clean-technology/ecologist-josh-donlan-on-bringing-sexy-animals-back-via- rewilding.html accessed 17 November 2014.

(38) C. Josh Donlan, Joel Berger, Carl E. Bock, et al., “Pleistocene Rewilding: An Optimistic Agenda for Twenty-First Century Conservation,” American Naturalist 168 (2006): 660–681, at 662–663.

(39) Donlan et al., “Pleistocene Rewilding,” 674.

(40) Sergey Zimov, “Pleistocene Park: Return of the Mammoth’s Ecosystem,” Science 308 (2005): 796–798.

(41) Emma Marris, “Reflecting the Past,” Nature 462 (2009): 30–32.

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(42) Joshua Tewkesbury and Haldre S. Rogers, “An Animal-Rich Future,” Science 345 (2014): 400.

(43) Marris, “Reflecting the Past”; “Oostvaardersplassen,” Natuurgebeiden Ontdek Nederland www.staatsbosbeheer.nl/Natuurgebeiden/Oostvaardersplassen/aspx, accessed 17 November 2014.

(44) “Oostvaardersplassen,” Nieuws & achtergronden, http://www.staatsbosbeheer.nl/Nieuws%20en%20achtergronden/Themas/Oostvaardersplassen.aspx; the Dutch site has more up-to-date information than the English one.: www.staasbosbeheer.nl/english.aspx accessed 17 November 2014.

(45) Elizabeth Kolbert, “Recall of the Wild,” The New Yorker, December 24, 2012; “Our Mission,” Rewilding Europe: Making Europe a Wilder Place, www.rewildingeurope.com/about/mission, accessed 17 November 2014.

(46) “Are Movies Science? Dinosaurs, Movies, and Reality,” DinoBuzz, n.d. http://www.ucmp.berkeley.edu/diapsids/buzz/popular.html accessed 17 November 2014.

(47) “India to Clone Cheetah,” BBC.com, 16 October 2000 http://news.bbc.co.uk/2/hi/south_asia/974858.stm, accessed 17 November 2014.

(48) Stewart Taggart, “Aussies Roaring on DNA Cloning,” Wired.com, 5 May 2000, http://archive.wired.com/science/discoveries/news/2000/05/36117, accessed 17 November 2014.

(49) Lesley Evans Ogden, “Extinction Is Forever … or Is It?” Bioscience 64 (2014), 469–475.

(50) Ogden, “Extinction Is Forever”.

(51) Ogden, “Extinction Is Forever”; see also Nathaniel Rich, “The Mammoth Cometh,” New York Times Magazine, March 2, 2014; Carl Zimmer, “Bringing Them Back to Life,” National Geographic, April 2013.

(52) See Rewilding Europe: Making Europe a Wilder Place, www.rewildingeurope.com, accessed 17 November 2014.

(53) Jacob Sherkow and Henry Greely, “What if Extinction Is Not Forever?” Science 340 (2013) 32–33; Carrie Friese and Claire Marris, “Making De-extinction Mundane?” PLOS Biology 12, no. 3 (March 2014): 1–3; on the ethical issues, Ben Minteer, “Is It Right to Reverse Extinction?” Nature 509 (2014): 261.

(54) C. Josh Donlan, “De-extinction in a Crisis Discipline,” Frontiers of Biogeography, 6 (2014): 25–28.

(55) George Monbiot, “Resurrecting Woolly Mammoths Is Exciting—But It’s a Fantasy,” Guardian, August 6, 2013.

(56) Sherkow and Greely, “What if Extinction?” 33.

Anita Guerrini Anita Guerrini, Oregon State University

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