hypothesis balance of nature

• What Is the Balance of Nature?

The ecosystem is often balanced when the living organisms such as plant, humans, and animals are in harmony. Humans are key in maintaining such a balance since the balance is dependent on their activities. However, people often carry out activities that are harmful and destructive to nature. Some, while interacting with nature, conserve it while others break it through the exploitation of forests, agricultural activities, and the introduction of invasive species. This destruction and conflict between humans and nature has led to the formation of protectionist groups and conservation activists. Because of the conflict between the nature and humans, there is a need for a balanced nature that will ensure not only the survival of plants and animals but also humans.

Natural Balance Theory

The balance of nature can be defined as a biological equilibrium between the living beings such as human, plants, and animals. At a stable equilibrium, the balance of nature asserts that any slight change in certain parameters will be corrected by a negative feedback which will eventually bring back the changed parameter to its original position of balance. The balance of nature applies in the case where there is interdependence in a population such as the predator-prey system or herbivores-vegetation system. The theory of permanently balanced nature has been criticized and dismissed by scientist, especially ecologists who have found that chaotic changes in populations are common. Despite the criticism, the theory is popular among the general public.

The concept that nature maintains its balance has existed for a long time. One of the most proponents of the theory was Herodotus who asserted that there is a perfect relationship between predators and preys which ensures that they remain in steady proportion to one another. In this wonderful relationship do not excessively feed on their prey. At some point, the theory of “balance of nature” dominated the ecological research and influence the management of the natural resources, leading to a popular doctrine among conservationist that nature would thrive if left to take care of itself and human intervention was unacceptable. The balance of nature concept was already in question by the beginning of the 20th century but the concept was abandoned by the scientists in the field of ecology in the last quarter of the century.

Human Intervention

The population of predators and prey often exhibit chaotic behaviors within the limit in which the population sizes in ways that may appear random and unpredictable but in real sense obeying the deterministic laws depending on the relationship between the population and the source of food as highlighted by Lotka-Volterra equation. Although humans have been accused of destroying the environment, some of their activities have contributed to creating modern day habitats. Some rainforests in Latin America were planted and transplanted by humans. The “fire-stick farming” practiced in Australian Aboriginal is an example of a human activity that modified the ecosystem.

The Fate of the Theory

Although the theory has been discredited by most of the ecologist, it is still widely held as true by the general public. In Midwestern America, the concept is widely held by both the science students and general students.

• Environment

More in Environment

Importance Of Wetlands

Meet 12 Incredible Conservation Heroes Saving Our Wildlife From Extinction

India's Leopard God, Waghoba, Aids Wildlife Conservation In The Country

Wildfires And Habitat Loss Are Killing Jaguars In The Amazon Rainforest

In India's Sundarbans: Where People Live Face-To-Face With Wild Tigers

Africa's "Thunderbird" Is At Risk Of Extinction

Why Is Biodiversity Critical To Life On Earth?

How Is Climate Change Impacting The Water Cycle?

An official website of the United States government

The .gov means it’s official. Federal government websites often end in .gov or .mil. Before sharing sensitive information, make sure you’re on a federal government site.

The site is secure. The https:// ensures that you are connecting to the official website and that any information you provide is encrypted and transmitted securely.

• Publications
• Account settings

Preview improvements coming to the PMC website in October 2024. Learn More or Try it out now .

• Advanced Search
• Journal List
• v.12(10); 2014 Oct

The “Balance of Nature”—Evolution of a Panchreston

Daniel simberloff.

Department of Ecology and Evolutionary Biology, University of Tennessee, Knoxville, Tennessee, United States

The notion of a “balance of nature” stretches back at least to the ancient Greeks, but with widely varying conceptions. Originally seen as bestowed by God to protect humankind, after Darwin, the balance came to be seen as fragile, arising by natural selection, and requiring human assistance for maintenance.

The earliest concept of a balance of nature in Western thought saw it as being provided by gods but requiring human aid or encouragement for its maintenance. With the rise of Greek natural philosophy, emphasis shifted to traits gods endowed species with at the outset, rather than human actions, as key to maintaining the balance. The dominance of a constantly intervening God in the Middle Ages lessened interest in the inherent features of nature that would contribute to balance, but the Reformation led to renewed focus on such features, particularly traits of species that would maintain all of them but permit none to dominate nature. Darwin conceived of nature in balance, and his emphasis on competition and frequent tales of felicitous species interactions supported the idea of a balance of nature. But Darwin radically changed its underlying basis, from God to natural selection. Wallace was perhaps the first to challenge the very notion of a balance of nature as an undefined entity whose accuracy could not be tested. His skepticism was taken up again in the 20th century, culminating in a widespread rejection of the idea of a balance of nature by academic ecologists, who focus rather on a dynamic, often chaotic nature buffeted by constant disturbances. The balance-of-nature metaphor, however, lives on in large segments of the public, representing a fragile aspect of nature and biodiversity that it is our duty to protect.

The notion of a “balance of nature” stretches back to early Greeks, who believed gods maintained it with the aid of human prayers, sacrifices, and rituals [1] . As Greek philosophers developed the idea of natural laws, human assistance in maintaining the balance did not disappear but was de-emphasized. Herodotus, for instance, the earliest known scholar to seek biological evidence for a balance of nature, asked how the different animal species each maintained their numbers, even though some species ate other species. Amassing facts and factoids, he saw divinely created predators' reproductive rates lower than those of prey, buttressing the idea of a providentially determined balance with a tale of a mutualism between Nile crocodiles beset with leeches and a plover species that feeds on them [1] . Two myths in Plato's Dialogues supported the idea of a balance of nature: the Timaeus myth, in which different elements of the universe, including living entities, are parts of a highly integrated “superorganism,” and the Protagoras myth, in which gods created each animal species with characteristics that would allow it to thrive and, having run out of biological traits, had to give man fire and superior intelligence [1] . Among Romans, Cicero followed Herodotus and Plato in advancing a balance of nature generated by different reproductive rates and traits among species, as well as interactions among species [1] .

The Middle Ages saw less interest in such pre-set devices as differential reproductive rates to keep nature in balance, perhaps because people believed in a God who would maintain the balance by frequent direct intervention [1] . The Reformation, however, fostered further development of the concept of a providential balance of nature set in motion at creation. Thomas Browne [2] added differential mortality rates to factors maintaining the balance, and Matthew Hale [3] proposed that lower rates of mortality for humans than for other animals maintain human dominance within a balanced nature and added vicissitudes of heat from the sun to the factors keeping any one species from getting out of hand.

The discovery of fossils that could not be ascribed to known living species severely challenged the idea of a God-given balance of nature, as they contradicted the idea of species divinely created with the necessary features for survival [4] . John Ray [5] suggested that the living representatives of such fossils would be found in unexplored parts of the earth, a solution that was viable until the great scientific explorations of the late 18th and early 19th centuries [4] . Ray also argued that what would now be termed different Grinnellian ecological niches demonstrated God's provision of each species with a space of its own in nature.

According to Egerton [1] , the earliest use of the term “balance” to refer specifically to ecology was probably by Ray's disciple, William Derham [6] , who asserted in 1714 that:

“The Balance of the Animal World is, throughout all Ages, kept even, and by a curious Harmony and just Proportion between the increase of all Animals, and the length of their Lives, the World is through all Ages well, but not over-stored.”

Derham recognized that human populations seemed to be endlessly increasing but saw this fact as a provision by God for future disasters. This explanation contrasts with that of Linnaeus [7] , who saw human and other populations endlessly increasing but believed the size of the earth was also increasing to accommodate them. Derham grappled with the issue of theodicy but failed to reconcile plagues of noxious animals with the balance of nature, seeing them rather as “Rods and Scourges to chastise us, as means to excite our Wisdom, Care, and Industry” [1] .

Derham's contemporary Richard Bradley [8] , [9] focused more on biological facts and less on Providence in sketching a more comprehensive account of an ecological balance of nature, taking account of the rapidly expanding knowledge of biodiversity, noting that each plant had its phytophagous insects, each insect its parasitic wasps or flies and predatory birds, concluding that “all Bodies have some Dependence upon one another; and that every distinct Part of Nature's Works is necessary for the Support of the rest; and that if any one was wanting, all the rest must consequently be out of Order.” Thus, he saw the balance as fragile rather than robust, in spite of a constantly intervening God. Linnaeus [10] similarly marshaled observations of species interactions to explain why no species increases to crowd out all others, adding competition to the predation, parasitism, and herbivory adduced by Bradley and also emphasizing the different roles (we might now say “niches”) of different species as allowing them all to coexist in a sort of superorganismic, balanced whole.

Unlike Derham, Georges-Louis Leclerc, Comte de Buffon [11] managed to reconcile animal plagues with a balanced nature. He perceived the balance of nature as dynamic, with all species fluctuating between relative rarity and abundance, so that whenever a species became overabundant, weather, predation, and competition for food would bring it back into balance. Buffon's successor as director of the Jardin des Plantes in Paris, Jacques-Henri Bernardin de Saint-Pierre [12] , was probably the first to associate ecological damage caused by biological invasions with a disruption of the balance of nature. Observing damage to introduced trees from insects accidentally introduced with them, he argued that failure to introduce the birds that would eat the insects led to the damage. William Paley [13] , perhaps the inspiration for today's advocates of “intelligent design,” analogized nature to a watch. One would assume a smoothly running watch was designed with purpose, and so too nature was designed by God with balance and a purpose.

In the 19th century, evolution burst on the scene, greatly influencing and ultimately modifying conceptions of a balance of nature. Fossils that seemed unrelated to any living species, as noted above, conflicted with the balance of nature, because they implied extinction, a manifestly unbalanced event that furthermore could be seen to imply that God had made a mistake. Whereas Ray had been able to argue that living exemplars of fossil species would be found in unexplored parts of the earth, by the 19th century, this explanation could be rejected. Jean-Baptiste Lamarck [14] resolved the conflict in a different way, arguing that species continually change, so the balance remains the same. The fossils thus represent ancestors of living species, not extinct lineages. Robert Chambers [15] , another early evolutionist, similarly saw fossils not as a paradox in a balanced nature but as a consequence of the fact that, as the physical environment changed, species either evolved or went extinct.

Alfred Russel Wallace was perhaps the first to question the very existence of a balance of nature, in a remarkable notebook entry, ca. 1855:

“Some species exclude all others in particular tracts. Where is the balance? When the locust devastates vast regions and causes the death of animals and man, what is the meaning of saying the balance is preserved… To human apprehension there is no balance but a struggle in which one often exterminates another” [16] .

In modern parlance, Wallace appears almost to be asking how “balance” could be defined in such a way that a balance of nature could be a testable hypothesis.

Darwin's theory of evolution by natural selection certainly explained the existence of fossils, and his emphasis on inevitable competition both between and within species downplayed the role of niche specialization propounded by Plato, Cicero, Linnaeus, Derham, and others [1] . Darwin nevertheless saw the ecological roles of the diversity of species as parts of an almost superorganismic nature, and his main contribution to the idea of a balance of nature was his constant emphasis on competition and other mortality factors that kept all species' populations in check [1] . His many metaphors and examples of the interactions among species, such as the tangled bank and the spinsters-cats-mice-bumblebees-clover stories in The Origin of Species [17] , contributed to a sense of a highly balanced nature, but one driven by natural selection constantly changing species, rather than by God either intervening or creating species with traits that ensure their continued existence. Unlike Wallace, Darwin did not raise the issue of whether nature was actually balanced and how we would know if it was not.

As ecology developed in the late 19th and early 20th centuries, it was inevitable that Wallace's question—how to define “balance”—would be raised again and that increasingly wide and quantitative study, especially at the population level, would be brought to bear on the matter. The work of the early dominant plant ecologist Frederic Clements and his followers, with Clements' notion of superorganismic communities [18] , provided at least tacit support for the idea of a balance of nature, but his contemporary Charles Elton [19] , a founder of the field of animal ecology and a leading student of animal population cycles, forcefully reprised Wallace's concern:

“‘The balance of nature’ does not exist, and perhaps never has existed. The numbers of wild animals are constantly varying to a greater or lesser extent, and the variations are usually irregular in period and always irregular in amplitude. Each variation in the numbers of one species causes direct and indirect repercussions on the numbers of the others, and since many of the latter are themselves independently varying in numbers, the resultant confusion is remarkable.”

Despite Elton's explicit skepticism, his depiction of energy flow through food chains and food webs was incorporated as a superorganismic analog to the physiology of individuals (e.g., [20] ). Henry Gleason, another critic of the superorganism concept, who depicted populations distributed independently, rather than in highly organized communities, was ignored at this time [21] .

However, beginning with three papers in Ecological Monographs in 1947, the superorganism concept was increasingly questioned and, within 25 years, Gleason was vindicated and his views largely accepted by ecologists [22] . During this same period, extensive work by population biologists again took up Elton's focus on population trajectories and contributed greatly to a growing recognition of the dynamism of nature and the fact that much of this dynamism did not seem regular or balanced [21] . The idea of a balanced nature did not immediately disappear among ecologists. For instance, a noteworthy book by C. B. Williams [23] , Patterns in the Balance of Nature , described the distribution of abundances within communities or regions as evincing statistical regularity that might be construed as a type of “balance of nature,” at least if changes in individual populations do not change certain statistical features (a hypothesis that Williams considered untested at the time). But the predominant view by ecologists of the 1960s saw the whole notion of a balance as, at best, irrelevant and, at worst, a distraction. Ehrlich and Birch [24] , for example, ridiculed the idea:

“The existence of supposed balance of nature is usually argued somewhat as follows. Species X has been in existence for thousands or perhaps millions of generations, and yet its numbers have never increased to infinity or decreased to zero. The same is true of the millions of other species still extant. During the next 100 years, the numbers of all these species will fluctuate; yet none will increase indefinitely, and only a few will become extinct… Such ‘observations’ are made the basis for the statement that population size is ‘controlled’ or ‘regulated,’ and that drastic changes in size are the results of upsetting the ‘balance of nature.’”

Another line of ecological research that became popular at the end of the 20th century was to equate “balance of nature” with some sort of equilibrium of numbers, usually of population sizes [25] , but sometimes of species richness. The problem remained that, with numbers that vary for whatever reason, it is still arbitrary just how much temporal variation can be accommodated within a process or phenomenon for it still to be termed equilibrial [26] . Often the decision on whether to perceive an ecological process as equilibrial seems to be based on whether there is some sort of homeostatic regulation of the numbers, such as density-dependence, which A. J. Nicholson [27] suggested as an argument against Elton's skepticism of the existence of a balance. The classic 1949 ecology text by Allee et al. [28] explicitly equated balance with equilibrium and cited various mechanisms, such as density-dependence, in support of its universality in nature [25] . Later similar sorts of mathematical arguments equated the mathematical stability of models representing nature with a balance of nature [29] , although the increasing recognition of stochastic aspects and chaotic mathematics of population fluctuations made it more difficult to perceive a balanced nature in population trajectories [21] .

For academic ecologists, the notion of a balance of nature has become passé, and the term is widely recognized as a panchreston [30] —a term that means so many different things to different people that it is useless as a theoretical framework or explanatory device. Much recent research has been devoted to emphasizing the dynamic aspects of nature and prominence of natural or anthropogenic disturbances, particularly as evidenced by vicissitudes of population sizes, and advances the idea that there is no such thing as a long-term equilibrium (e.g., [31] , [32] ). Some authors explicitly relate this research to a rejection of the concept of a balance of nature (e.g., [33] – [35] ), Pickett et al. [33] going so far as to say it must be replaced by a different metaphor, the “flux of nature.”

The issue is confounded by the fact that the perception of balance can be sought at different levels (populations, communities, ecosystems) and spatial scales. Much of the earlier discussion of a balance was at the population and community levels—Browne, Hale, Bradley, Linnaeus, Buffon, Bernardin de Saint-Pierre, and Darwin saw balance in the limited fluctuations of populations and the interactions of populations as one force imposing the limits. The proponents of density-dependent population regulation fall in this category as well [36] , [37] . As a balance is sought at the community and ecosystem levels, the sorts of evidence brought to bear on the matter become more complicated and abstract [37] , [38] . It is increasingly difficult to imagine what sorts of empirical or observational data could test the notion of a balance. For instance, Williams's balance of nature—evidenced by a particular statistical distribution of population sizes—would not be perceived as balanced by many observers in light of the fact that entire populations can crash, explode, or even go extinct within the constraint of a statistical distribution of a given shape. Early claims of a balance at the highest level, such as the various superorganisms (Plato's Timaeus myth, Paley's watch metaphor, Clements's superorganismic plant community) can hardly be seen as anything other than metaphors rather than testable hypotheses and have fallen from favor. The most expansive conception of a balance of nature—the Gaia hypothesis [39] —has been almost universally rejected by scientists [40] . The advent and growing acceptance of the metapopulation concept of nature [41] also complicates the search for balance in bounded population fluctuations. Spatially limited individual populations can arise, fluctuate wildly, and even go extinct, while suitable dynamics maintain the widespread metapopulation as a whole.

Yet, the idea of a balance of nature lives on in the popular imagination, especially among conservationists and environmentalists. However, the usual use of the metaphor in an environmental context suggests that the balance, whether given by God or produced by evolution, is a fragile balance, one that needs human actions for its maintenance. Through the 18th century, the balance of nature was probably primarily a comforting construct—it would protect us; it represented some sort of benign governance in the face of occasional awful events. When Darwin replaced God as the determinant of the balance with natural selection, the comfort of a balance of nature was not so overarching, if there was any comfort at all. Today, ecologists do not even recognize a balance, and those members of the public who do, see it as something we must protect if we are ever to reap benefits from it in the future (e.g., wetlands that might help ameliorate flooding from storms and sea-level rise). This shift is clear in the writings of Bill McKibben [42] , [43] , who talks frequently about balance, but about balance with nature, not balance of nature, and how humankind is headed towards a catastrophic future if it does not act promptly and radically to rebalance society with nature.

Acknowledgments

My debt will be obvious to a remarkable paper by Frank N. Edgerton on the history of the concept of a balance of nature.

Funding Statement

No funding was received for this work.

Nature is a highly dynamic system, experts say, and there's no such thing as perfect balance.

• ENVIRONMENT

The ‘balance of nature’ is an enduring concept. But it’s wrong.

From the ancient Greeks to the Lion King, people have sought balance in nature—but the real world isn’t like that.

Strolling across his animated kingdom—Pride Rock in the distance—Mufasa explains to his young son Simba : “Everything you see exists together, in a delicate balance.”

The line is a hallmark of the Disney movie the Lion King , which debuted in 1994. A visually rich update of the classic, also by Disney (the Walt Disney Company is majority owner of National Geographic Partners), hit theaters this July. In the quarter-century gap, the film saw a few significant changes—hand-drawn animation gave way to life-like computer-generated graphics and Beyonce’s character Nala got a whole new song. But other aspects of the movie remained unchanged, including Mufasa’s original lesson about nature being in balance.

John Kricher just rewatched the scene from 1994. While the Wheaton College professor and author of The Balance of Nature: Ecology's Enduring Myth is a fan of James Earl Jones’ deep baritone delivery, he says “it’s not sound science.”

Scientists have long abandoned the idea of there being a “balance of nature,” in favor of more dynamic ecological frameworks. But, having been ingrained in popular culture over millennia, it’s proven much harder for the public to shake. The metaphor is alive and well today, appearing everywhere from newspapers , Legos , and a health food brand name to social media , and, of course, the Lion King reboot.

The misconception impacts everything from conservation management to climate change policy; and it’s a concept that scientists would like to see plucked from the public’s vocabulary. “It's a satisfying term,” says Kricher. “But it's not useful.”

From Herodotus to Disney

The notion that nature exists in some sort of balance, or harmony, dates back to at least the ancient Greeks . The Greek writer Herodotus, for instance, was fascinated by the apparently balanced relationship between predator and prey. Cicero, a Roman politician and philosopher, imbued the idea with religious significance, presenting it as evidence of the wisdom of a Creator.

“It’s so old,” said Kim Cuddington , a professor at the University of Waterloo, in Canada, that “it's very much embedded, at least in Western culture.”

Even among scientists, the balance of nature concept persisted for centuries. Charles Darwin, the famous naturalist, alluded to it in his work on natural selection, as did his contemporaries, such as Herbert Spencer. The approach bled into the twentieth century, with, for example, the belief that leaving the wilderness wild was the best method of conservation or that pollution was a disruptor of the natural order.

But it was around that same time that people also began to challenge balance of nature thinking. In 1949, environmentalist Aldo Leopold wrote, “The image commonly employed in conservation education is 'the balance of nature.' For reasons too lengthy to detail here, this figure of speech fails to describe accurately what little we know about the land mechanism.”

Stunning photos of the Earth

Also at that time, science was becoming more data driven, and ecology a more established discipline. “When the data don't support it, then you have to revise your idea,” said Kricher, explaining that that’s exactly what happened with the balance of nature.

Ecologists shifted away from community-based sociological models to increasingly mathematical, individualist theories. And, throughout the 1970s and 1980s, the phrase balance of nature largely disappeared from the scientific lexicon . “Ecologists,” said Kricher, “had a tacit understanding that the [phrase] was largely metaphorical.”

The public, however, still employs the phrase liberally. The expression is often used one of two ways, said Cuddington. Sometimes the balance is depicted as fragile, delicate, and easily disturbed. Other times it’s the opposite—that the balance of nature is so powerful that it can correct any imbalances on its own. According to Cuddington, “they’re both wrong.”

Constant change

In the 1950s, Robert MacArthur went into the remote woods of coastal Maine to observe warbler populations. He found that five different species coexisted by utilizing different parts of the same tree. The phenomenon, known as niche partitioning, implied an equilibrium. A balance.

You May Also Like

Keep mosquitoes away with these simple tips, backed by science

14 bison came to Catalina for a movie. 100 years later, what’s next?

These breathtaking natural wonders no longer exist

The study became a foundation of ecology—one that most students read in class. That included Bik Wheeler, who first came across the work while pursuing his master’s degree. Realizing that he lived near MacArthur’s field site, he proposed re-creating the seminal research.

That was 2014. In the more than half-century that had passed since MacArthur’s original research, the forest he had worked in had largely gone untouched. So, aside from a few technological improvements—such as using lasers instead of stopwatches—Wheeler was able to retrace MacArthur’s steps. His results, however, were different.

Wheeler’s paper is still pending publication and peer review, so he was unable to talk about specifics. But he says he observed only two of the same species as MacArthur , as well as a few new ones.

In other words, “it's a dynamic system,” he said. “It's not static.”

The changes that Wheeler saw are emblematic of the broader shift that’s happened across ecology, and has had real-life implications. Both the delicate and stalwart interpretations of “balance” imply that nature should be left to its own devices; that human interference ought to be minimal.

The updated view is that “change is constant,” said Matt Palmer, an ecologist at Columbia University. And as the new approach took hold, conservation and management policies also adapted. “In some ways it argues for a stronger hand in managing ecosystems or natural resources,” he said. “It's going to take human intervention.”

Palmer pointed to assisted migration—in which people help animals move across obstacles such as roads or those caused by the changing climate—as one common conservation technique that bucks the notion of a natural balance. The reintroduction of wolves in places such as Yellowstone is another example. Others noted controlled-burn fires in forest management.

But the most obvious, and pressing, manifestation is the looming climate crisis, says Corinne Zimmerman , a psychologist at Illinois State University. While the vast majority of scientists agree that efforts to address climate change must involve human action , a public misconception about nature being in balance could inhibit progress. “If nature is all robust and fine, she'll take care of herself, we don't have to do anything about our carbon footprint,” she said. “It's a very naive understanding of nature.”

Still, the term persists. A 2007 study by Zimmerman and Cuddington found that even being exposed to scientific evidence against the balance of nature did little to change people’s opinions. Those surveyed, Cuddington told Pacific Standard magazine at the time, were “almost unable to reason logically about environmental problems because they keep bumping into this cultural concept.”

The flux of nature?

Dislodging the balance of nature from the popular psyche won’t be easy. Back in the 1980s, ecologist Steward Pickett tried to aid that process by proposing a replacement: the “flux of nature.”

“I thought that there needed to be some short-hand alternative,” he said.

Although the flux alternative hasn’t yet caught on, Pickett still thinks it’s a more useful framing. The balance of nature, he says, has left the scientific discourse because it’s “vague and loaded” and the public ought to follow suit.

“We have to accept responsibility for what we're doing,” he said, “and not just say that nature will take care of it.”

Related Topics

• CLIMATE CHANGE
• ENVIRONMENT AND CONSERVATION

Extreme heat is the future. Here are 10 practical ways to manage it.

Another weapon to fight climate change? Put carbon back where we found it

Rats invaded paradise. Here’s how paradise fought back.

For Antarctica’s emperor penguins, ‘there is no time left’

Listen to 30 years of climate change transformed into haunting music

• Environment
• Paid Content

History & Culture

• History & Culture
• Mind, Body, Wonder
• Terms of Use
• Privacy Policy
• Your US State Privacy Rights
• Children's Online Privacy Policy
• Interest-Based Ads
• About Nielsen Measurement
• Do Not Sell or Share My Personal Information
• Nat Geo Home
• Attend a Live Event
• Book a Trip
• Inspire Your Kids
• Shop Nat Geo
• Visit the D.C. Museum
• Learn About Our Impact
• Support Our Mission
• Advertise With Us
• Customer Service
• Renew Subscription
• Manage Your Subscription
• Work at Nat Geo
• Sign Up for Our Newsletters
• Contribute to Protect the Planet

Copyright © 1996-2015 National Geographic Society Copyright © 2015-2024 National Geographic Partners, LLC. All rights reserved

• A-Z Publications

Annual Review of Marine Science

Volume 14, 2022, review article, the balance of nature: a global marine perspective.

• Constantin W. Arnscheidt 1 , and Daniel H. Rothman 1
• View Affiliations Hide Affiliations Affiliations: Lorenz Center, Department of Earth, Atmospheric, and Planetary Sciences, Massachusetts Institute of Technology, Cambridge, Massachusetts 02139, USA; email: [email protected] [email protected]
• Vol. 14:49-73 (Volume publication date January 2022) https://doi.org/10.1146/annurev-marine-010318-095212
• First published as a Review in Advance on June 11, 2021
• Copyright © 2022 by Annual Reviews. All rights reserved

The ancient idea of the balance of nature continues to influence modern perspectives on global environmental change. Assumptions of stable biogeochemical steady states and linear responses to perturbation are widely employed in the interpretation of geochemical records. Here, we review the dynamics of the marine carbon cycle and its interactions with climate and life over geologic time, focusing on what the record of past changes can teach us about stability and instability in the Earth system. Emerging themes include the role of amplifying feedbacks in producing past carbon cycle disruptions, the importance of critical rates of change in the context of mass extinctions and potential Earth system tipping points, and the application of these ideas to the modern unbalanced carbon cycle. A comprehensive dynamical understanding of the marine record of global environmental disruption will be of great value in understanding the long-term consequences of anthropogenic change.

Article metrics loading...

Full text loading...

Literature Cited

• Alexandrov DV , Bashkirtseva IA , Crucifix M , Ryashko LB. 2021 . Nonlinear climate dynamics: from deterministic behavior to stochastic excitability and chaos. Phys. Rep. 902 : 1– 60 [Google Scholar]
• Alroy J. 2008 . Dynamics of origination and extinction in the marine fossil record. PNAS 105 : Suppl. 1 11536– 42 [Google Scholar]
• Alroy J. 2014 . Accurate and precise estimates of origination and extinction rates. Paleobiology 40 : 374– 97 [Google Scholar]
• Alvarez LW , Alvarez W , Asaro F , Michel HV. 1980 . Extraterrestrial cause for the Cretaceous-Tertiary extinction. Science 208 : 1095– 108 [Google Scholar]
• Archer D. 2005 . Fate of fossil fuel CO 2 in geologic time. J. Geophys. Res. Oceans 110 : C09S05 [Google Scholar]
• Archer D. 2010 . The Global Carbon Cycle Princeton, NJ: Princeton Univ. Press [Google Scholar]
• Archer D , Kheshgi H , Maier-Reimer E. 1998 . Dynamics of fossil fuel CO 2 neutralization by marine CaCO 3 . Glob. Biogeochem. Cycles 12 : 259– 76 [Google Scholar]
• Archer D , Winguth A , Lea D , Mahowald N. 2000 . What caused the glacial/interglacial atmospheric p CO 2 cycles?. Rev. Geophys. 38 : 159– 89 [Google Scholar]
• Arnscheidt CW , Rothman DH. 2020 . Routes to global glaciation. Proc. R. Soc. A 476 : 20200303 [Google Scholar]
• Arrhenius S. 1896 . On the influence of carbonic acid in the air upon the temperature of the ground. Lond. Edinb. Dublin Philos. Mag. J. Sci. 41 : 237– 76 [Google Scholar]
• Ashwin P , Wieczorek S , Vitolo R , Cox P. 2012 . Tipping points in open systems: bifurcation, noise-induced and rate-dependent examples in the climate system. Philos. Trans. R. Soc. A 370 : 1166– 84 [Google Scholar]
• Bak P. 1996 . How Nature Works: The Science of Self-Organized Criticality New York: Springer [Google Scholar]
• Bambach RK. 2006 . Phanerozoic biodiversity mass extinctions. Annu. Rev. Earth Planet. Sci. 34 : 127– 55 [Google Scholar]
• Barnosky AD , Matzke N , Tomiya S , Wogan G , Swartz B et al. 2011 . Has the Earth's sixth mass extinction already arrived?. Nature 471 : 51– 57 [Google Scholar]
• Berner RA. 1992 . Weathering, plants, and the long-term carbon cycle. Geochim. Cosmochim. Acta 56 : 3225– 31 [Google Scholar]
• Berner RA. 2004 . The Phanerozoic Carbon Cycle: CO 2 and O 2 New York: Oxford Univ. Press [Google Scholar]
• Berner RA , Caldeira K. 1997 . The need for mass balance and feedback in the geochemical carbon cycle. Geology 25 : 955– 56 [Google Scholar]
• Berner RA , Lasaga AC , Garrels RM. 1983 . The carbonate-silicate geochemical cycle and its effect on atmospheric carbon dioxide over the past 100 million years. Am. J. Sci. 283 : 641– 83 [Google Scholar]
• Blackburn TJ , Olsen PE , Bowring SA , McLean NM , Kent DV et al. 2013 . Zircon U-Pb geochronology links the end-Triassic extinction with the Central Atlantic Magmatic Province. Science 340 : 941– 45 [Google Scholar]
• Brand U , Blamey N , Garbelli C , Griesshaber E , Posenato R et al. 2016 . Methane hydrate: killer cause of Earth's greatest mass extinction. Palaeoworld 25 : 496– 507 [Google Scholar]
• Broecker WS. 1982 . Glacial to interglacial changes in ocean chemistry. Prog. Oceanogr. 11 : 151– 97 [Google Scholar]
• Broecker WS , Peng TH. 1987 . The role of CaCO 3 compensation in the glacial to interglacial atmospheric CO 2 change. Glob. Biogeochem. Cycles 1 : 15– 29 [Google Scholar]
• Burgess SD , Bowring SA. 2015 . High-precision geochronology confirms voluminous magmatism before, during, and after Earth's most severe extinction. Sci. Adv. 1 : e1500470 [Google Scholar]
• Caldeira K. 1991 . Continental-pelagic carbonate partitioning and the global carbonate-silicate cycle. Geology 19 : 204– 6 [Google Scholar]
• Ceballos G , Ehrlich PR , Barnosky AD , Garca A , Pringle RM , Palmer TM. 2015 . Accelerated modern human–induced species losses: entering the sixth mass extinction. Sci. Adv. 1 : e1400253 [Google Scholar]
• Ciais P , Sabine C , Bala G , Bopp L , Brovkin V et al. 2013 . Carbon and other biogeochemical cycles. Climate Change 2013: The Physical Science Basis; Contribution of Working Group I to the Fifth Assessment Report of the Intergovernmental Panel on Climate Change TF Stocker, D Qin, G-K Plattner, M Tignor, SK Allen et al. 465– 570 Cambridge, UK: Cambridge Univ. Press [Google Scholar]
• Clapham ME , Renne PR. 2019 . Flood basalts and mass extinctions. Annu. Rev. Earth Planet. Sci. 47 : 275– 303 [Google Scholar]
• Colbourn G , Ridgwell A , Lenton T. 2015 . The time scale of the silicate weathering negative feedback on atmospheric CO 2 . Glob. Biogeochem. Cycles 29 : 583– 96 [Google Scholar]
• Courtillot V , Fluteau F. 2014 . A review of the embedded time scales of flood basalt volcanism with special emphasis on dramatically short magmatic pulses. Volcanism, Impacts, and Mass Extinctions: Causes and Effects G Keller, AC Kerr Boulder, CO: Geol. Soc. Am. https://doi.org/10.1130/2014.2505(15) [Crossref] [Google Scholar]
• Cramer B , Jarvis I 2020 . Carbon isotope stratigraphy. Geologic Time Scale 2020 FM Gradstein, JG Ogg, M Schmitz, G Ogg 309– 43 Amsterdam: Elsevier [Google Scholar]
• Crucifix M. 2012 . Oscillators and relaxation phenomena in Pleistocene climate theory. Philos. Trans. R. Soc. A 370 : 1140– 65 [Google Scholar]
• Cui H , Kaufman AJ , Xiao S , Zhou C , Liu XM. 2017 . Was the Ediacaran Shuram Excursion a globally synchronized early diagenetic event? Insights from methane-derived authigenic carbonates in the uppermost Doushantuo Formation, South China. Chem. Geol. 450 : 59– 80 [Google Scholar]
• Cuvier G. 1812 (1997 . Preliminary discourse. Translated in Georges Cuvier, Fossil Bones, and Geological Catastrophes: New Translations and Interpretations of the Primary Texts ed. MJS Rudwick 183– 252 Chicago: Univ. Chicago Press [Google Scholar]
• D'Antonio MP , Ibarra DE , Boyce CK. 2020 . Land plant evolution decreased, rather than increased, weathering rates. Geology 48 : 29– 33 [Google Scholar]
• Darwin C 1859 . On the Origin of Species London: Murray [Google Scholar]
• Davis Barnes B , Husson JM , Peters SE 2020 . Authigenic carbonate burial in the Late Devonian–Early Mississippian Bakken Formation (Williston Basin, USA). Sedimentology 67 : 2065– 94 [Google Scholar]
• DeConto RM , Galeotti S , Pagani M , Tracy D , Schaefer K et al. 2012 . Past extreme warming events linked to massive carbon release from thawing permafrost. Nature 484 : 87– 91 [Google Scholar]
• Derham W. 1713 . Physico-Theology; or, A Demonstration of the Being and Attributes of God, from His Works of Creation London: Innys [Google Scholar]
• Dickens GR. 2003 . Rethinking the global carbon cycle with a large, dynamic and microbially mediated gas hydrate capacitor. Earth Planet. Sci. Lett. 213 : 169– 83 [Google Scholar]
• Dijkstra HA. 2013 . Nonlinear Climate Dynamics Cambridge, UK: Cambridge Univ. Press [Google Scholar]
• Dunkley Jones T , Ridgwell A , Lunt D , Maslin M , Schmidt D , Valdes P 2010 . A Palaeogene perspective on climate sensitivity and methane hydrate instability. Philos. Trans. R. Soc. A 368 : 2395– 415 [Google Scholar]
• Dyer B , Maloof AC , Higgins JA. 2015 . Glacioeustasy, meteoric diagenesis, and the carbon cycle during the Middle Carboniferous. Geochem. Geophys. Geosyst. 16 : 3383– 99 [Google Scholar]
• Edmond JM , Huh Y. 2003 . Non-steady state carbonate recycling and implications for the evolution of atmospheric . Earth Planet. Sci. Lett. 216 : 125– 39 [Google Scholar]
• Egerton FN. 1973 . Changing concepts of the balance of nature. Q. Rev. Biol. 48 : 322– 50 [Google Scholar]
• Emerson SR , Hedges JI. 2008 . Chemical Oceanography and the Marine Carbon Cycle Cambridge, UK: Cambridge Univ. Press [Google Scholar]
• Emiliani C. 1955 . Pleistocene temperatures. J. Geol. 63 : 538– 78 [Google Scholar]
• Fantle MS , Barnes BD , Lau KV. 2020 . The role of diagenesis in shaping the geochemistry of the marine carbonate record. Annu. Rev. Earth Planet. Sci. 48 : 549– 83 [Google Scholar]
• Feulner G. 2012 . The faint young Sun problem. Rev. Geophys. 50 : RG2006 [Google Scholar]
• Foote E. 1856 . Circumstances affecting the heat of the sun's rays. Am. J. Sci. Arts 22 : 382– 83 [Google Scholar]
• Foster GL , Royer DL , Lunt DJ. 2017 . Future climate forcing potentially without precedent in the last 420 million years. Nat. Commun. 8 : 14845 [Google Scholar]
• Ghil M , Childress S. 1987 . Topics in Geophysical Fluid Dynamics: Atmospheric Dynamics, Dynamo Theory, and Climate Dynamics New York: Springer [Google Scholar]
• Gildor H , Tziperman E. 2000 . Sea ice as the glacial cycles' climate switch: role of seasonal and orbital forcing. Paleoceanography 15 : 605– 15 [Google Scholar]
• Grossman E , Joachimski M 2020 . Oxygen isotope stratigraphy. Geologic Time Scale 2020 FM Gradstein, JG Ogg, M Schmitz, G Ogg 279– 307 Amsterdam: Elsevier [Google Scholar]
• Gutjahr M , Ridgwell A , Sexton PF , Anagnostou E , Pearson PN et al. 2017 . Very large release of mostly volcanic carbon during the Palaeocene–Eocene Thermal Maximum. Nature 548 : 573– 77 [Google Scholar]
• Hallam A , Wignall PB. 1997 . Mass Extinctions and Their Aftermath Oxford, UK: Oxford Univ. Press [Google Scholar]
• Hayes JM , Strauss H , Kaufman AJ. 1999 . The abundance of 13 C in marine organic matter and isotopic fractionation in the global biogeochemical cycle of carbon during the past 800 Ma. Chem. Geol. 161 : 103– 25 [Google Scholar]
• Hays JD , Imbrie J , Shackleton NJ. 1976 . Variations in the Earth's orbit: pacemaker of the ice ages. Science 194 : 1121– 32 [Google Scholar]
• Higgins JA , Blättler C , Lundstrom E , Santiago-Ramos D , Akhtar A et al. 2018 . Mineralogy, early marine diagenesis, and the chemistry of shallow-water carbonate sediments. Geochim. Cosmochim. Acta 220 : 512– 34 [Google Scholar]
• Hofmann M , Schellnhuber HJ 2009 . Oceanic acidification affects marine carbon pump and triggers extended marine oxygen holes. PNAS 106 : 3017– 22 [Google Scholar]
• Husson JM , Higgins JA , Maloof AC , Schoene B. 2015 . Ca and Mg isotope constraints on the origin of Earth's deepest δ 13 C excursion. Geochim. Cosmochim. Acta 160 : 243– 66 [Google Scholar]
• Husson JM , Linzmeier BJ , Kitajima K , Ishida A , Maloof AC et al. 2020 . Large isotopic variability at the micron-scale in ‘Shuram’ excursion carbonates from South Australia. Earth Planet. Sci. Lett. 538 : 116211 [Google Scholar]
• Izhikevich EM. 2007 . Dynamical Systems in Neuroscience: The Geometry of Excitability and Bursting Cambridge, MA: MIT Press [Google Scholar]
• Joachimski MM , Lai X , Shen S , Jiang H , Luo G et al. 2012 . Climate warming in the latest Permian and the Permian–Triassic mass extinction. Geology 40 : 195– 98 [Google Scholar]
• Kirchner JW. 1989 . The Gaia hypothesis: Can it be tested?. Rev. Geophys. 27 : 223– 35 [Google Scholar]
• Kirchner JW. 2002 . The Gaia hypothesis: fact, theory, and wishful thinking. Clim. Change 52 : 391– 408 [Google Scholar]
• Kirtland Turner S 2018 . Constraints on the onset duration of the Paleocene–Eocene Thermal Maximum. Philos. Trans. R. Soc. A 376 : 20170082 [Google Scholar]
• Kirtland Turner S , Sexton PF , Charles CD , Norris RD 2014 . Persistence of carbon release events through the peak of early Eocene global warmth. Nat. Geosci. 7 : 748– 51 [Google Scholar]
• Knoll AH , Bambach RK , Payne JL , Pruss S , Fischer WW. 2007 . Paleophysiology and end-Permian mass extinction. Earth Planet. Sci. Lett. 256 : 295– 313 [Google Scholar]
• Kolbert E. 2014 . The Sixth Extinction: An Unnatural History London: A&C Black [Google Scholar]
• Kump LR , Arthur MA. 1999 . Interpreting carbon-isotope excursions: carbonates and organic matter. Chem. Geol. 161 : 181– 98 [Google Scholar]
• Kurtz A , Kump L , Arthur M , Zachos J , Paytan A. 2003 . Early Cenozoic decoupling of the global carbon and sulfur cycles. Paleoceanography 18 : 1090 [Google Scholar]
• Le Treut H , Ghil M 1983 . Orbital forcing, climatic interactions, and glaciation cycles. J. Geophys. Res. Oceans 88 : 5167– 90 [Google Scholar]
• Lenton TM , Daines SJ , Dyke JG , Nicholson AE , Wilkinson DM , Williams HT. 2018 . Selection for Gaia across multiple scales. Trends Ecol. Evol. 33 : 633– 45 [Google Scholar]
• Lenton TM , Held H , Kriegler E , Hall JW , Lucht W et al. 2008 . Tipping elements in the Earth's climate system. PNAS 105 : 1786– 93 [Google Scholar]
• Lisiecki LE , Raymo ME. 2005 . A Pliocene-Pleistocene stack of 57 globally distributed benthic δ 18 O records. Paleoceanography 20 : PA1003 [Google Scholar]
• Lotka AJ. 1925 . Elements of Physical Biology Baltimore, MD: Williams & Wilkins [Google Scholar]
• Lourens LJ , Sluijs A , Kroon D , Zachos JC , Thomas E et al. 2005 . Astronomical pacing of late Palaeocene to early Eocene global warming events. Nature 435 : 1083– 87 [Google Scholar]
• Lovejoy AO. 1936 . The Great Chain of Being Cambridge, MA: Harvard Univ. Press [Google Scholar]
• Lovejoy S. 2015 . A voyage through scales, a missing quadrillion and why the climate is not what you expect. Clim. Dyn. 44 : 3187– 210 [Google Scholar]
• Lovelock JE , Margulis L. 1974 . Atmospheric homeostasis by and for the biosphere: the Gaia hypothesis. Tellus 26 : 2– 10 [Google Scholar]
• Lunt DJ , Ridgwell A , Sluijs A , Zachos J , Hunter S , Haywood A. 2011 . A model for orbital pacing of methane hydrate destabilization during the Palaeogene. Nat. Geosci. 4 : 775– 78 [Google Scholar]
• Lyell C. 1832 . Principles of Geology, Being an Attempt to Explain the Former Changes of the Earth's Surface, by Reference to Causes Now in Operation , Vol. 2 London: Murray [Google Scholar]
• Macdonald FA , Swanson-Hysell NL , Park Y , Lisiecki L , Jagoutz O. 2019 . Arc-continent collisions in the tropics set Earth's climate state. Science 364 : 181– 84 [Google Scholar]
• Maher K , Chamberlain C. 2014 . Hydrologic regulation of chemical weathering and the geologic carbon cycle. Science 343 : 1502– 4 [Google Scholar]
• May RM. 1973 . Stability and Complexity in Model Ecosystems Princeton, NJ: Princeton Univ. Press [Google Scholar]
• Mayr E. 1982 . The Growth of Biological Thought: Diversity, Evolution, and Inheritance Cambridge, MA: Harvard Univ. Press [Google Scholar]
• McInerney FA , Wing SL. 2011 . The Paleocene-Eocene Thermal Maximum: a perturbation of carbon cycle, climate, and biosphere with implications for the future. Annu. Rev. Earth Planet. Sci. 39 : 489– 516 [Google Scholar]
• Milanković M. 1941 . Kanon der Erdbestrahlung und seine Anwendung auf das Eiszeitenproblem Belgrade: K. Serb. Akad. [Google Scholar]
• Mitnick EH , Lammers LN , Zhang S , Zaretskiy Y , DePaolo DJ. 2018 . Authigenic carbonate formation rates in marine sediments and implications for the marine δ 13 C record. Earth Planet. Sci. Lett. 495 : 135– 45 [Google Scholar]
• Newell ND. 1963 . Crises in the history of life. Scientific American Feb., pp 76– 95 [Google Scholar]
• Paillard D , Parrenin F. 2004 . The Antarctic ice sheet and the triggering of deglaciations. Earth Planet. Sci. Lett. 227 : 263– 71 [Google Scholar]
• Payne JL , Clapham ME. 2012 . End-Permian mass extinction in the oceans: an ancient analog for the twenty-first century?. Annu. Rev. Earth Planet. Sci. 40 : 89– 111 [Google Scholar]
• Petit JR , Jouzel J , Raynaud D , Barkov NI , Barnola JM et al. 1999 . Climate and atmospheric history of the past 420,000 years from the Vostok ice core, Antarctica. Nature 399 : 429– 36 [Google Scholar]
• Pierrehumbert RT. 2010 . Principles of Planetary Climate Cambridge, UK: Cambridge Univ. Press [Google Scholar]
• Pope A. 1733 . An Essay on Man: In Epistles to a Friend Part I London: Wilford. , 2nd ed.. [Google Scholar]
• Ridgwell A. 2005 . A Mid Mesozoic Revolution in the regulation of ocean chemistry. Mar. Geol. 217 : 339– 57 [Google Scholar]
• Ridgwell A , Edwards U 2007 . Geological carbon sinks. Greenhouse Gas Sinks D Reay, CN Hewitt, K Smith, J Grace 74– 97 Wallingford, UK: CABI [Google Scholar]
• Ridgwell A , Hargreaves J. 2007 . Regulation of atmospheric CO 2 by deep-sea sediments in an Earth system model. Glob. Biogeochem. Cycles 21 : GB2008 [Google Scholar]
• Ridgwell A , Zeebe RE. 2005 . The role of the global carbonate cycle in the regulation and evolution of the Earth system. Earth Planet. Sci. Lett. 234 : 299– 315 [Google Scholar]
• Riebesell U , Körtzinger A , Oschlies A 2009 . Sensitivities of marine carbon fluxes to ocean change. PNAS 106 : 20602– 9 [Google Scholar]
• Rothman DH. 2015 . Earth's carbon cycle: a mathematical perspective. Bull. Am. Math. Soc. 52 : 47– 64 [Google Scholar]
• Rothman DH. 2017 . Thresholds of catastrophe in the Earth system. Sci. Adv. 3 : e1700906 [Google Scholar]
• Rothman DH. 2019 . Characteristic disruptions of an excitable carbon cycle. PNAS 116 : 14813– 22 [Google Scholar]
• Rothman DH , Fournier GP , French KL , Alm EJ , Boyle EA et al. 2014 . Methanogenic burst in the end-Permian carbon cycle. PNAS 111 : 5462– 67 [Google Scholar]
• Rothman DH , Hayes JM , Summons RE 2003 . Dynamics of the Neoproterozoic carbon cycle. PNAS 100 : 8124– 29 [Google Scholar]
• Rudwick MJS. 2014 . Earth's Deep History: How It Was Discovered and Why It Matters Chicago: Univ. Chicago Press [Google Scholar]
• Sagan C , Mullen G. 1972 . Earth and Mars: evolution of atmospheres and surface temperatures. Science 177 : 52– 56 [Google Scholar]
• Saltzman B. 2002 . Dynamical Paleoclimatology: Generalized Theory of Global Climate Change San Diego, CA: Academic [Google Scholar]
• Saltzman B , Hansen AR , Maasch KA. 1984 . The late Quaternary glaciations as the response of a three-component feedback system to Earth-orbital forcing. J. Atmos. Sci. 41 : 3380– 89 [Google Scholar]
• Sarmiento JL , Gruber N. 2006 . Ocean Biogeochemical Dynamics Princeton, NJ: Princeton Univ. Press [Google Scholar]
• Schoene B , Eddy MP , Samperton KM , Keller CB , Keller G et al. 2019 . U-Pb constraints on pulsed eruption of the Deccan Traps across the end-Cretaceous mass extinction. Science 363 : 862– 66 [Google Scholar]
• Schrag DP , Higgins JA , Macdonald FA , Johnston DT. 2013 . Authigenic carbonate and the history of the global carbon cycle. Science 339 : 540– 43 [Google Scholar]
• Self S , Thordarson T , Widdowson M. 2005 . Gas fluxes from flood basalt eruptions. Elements 1 : 283– 87 [Google Scholar]
• Sepkoski D. 2020 . Catastrophic Thinking: Extinction and the Value of Diversity from Darwin to the Anthropocene Chicago: Univ. Chicago Press [Google Scholar]
• Sexton PF , Norris RD , Wilson PA , Pälike H , Westerhold T et al. 2011 . Eocene global warming events driven by ventilation of oceanic dissolved organic carbon. Nature 471 : 349– 52 [Google Scholar]
• Shackleton NJ 1977 . Carbon-13 in Uvigerina: tropical rainforest history and the equatorial Pacific carbonate dissolution cycles. The Fate of Fossil Fuel CO 2 in the Oceans NR Andersen, A Malahoff 401– 27 New York: Plenum [Google Scholar]
• Sherwood S , Webb MJ , Annan JD , Armour K , Forster PM et al. 2020 . An assessment of Earth's climate sensitivity using multiple lines of evidence. Rev. Geophys. 58 : e2019RG000678 [Google Scholar]
• Sigman DM , Boyle EA. 2000 . Glacial/interglacial variations in atmospheric carbon dioxide. Nature 407 : 859– 69 [Google Scholar]
• Simberloff D. 2014 . The “balance of nature”—evolution of a panchreston. PLOS Biol 12 : e1001963 [Google Scholar]
• Sprain CJ , Renne PR , Vanderkluysen L , Pande K , Self S , Mittal T. 2019 . The eruptive tempo of Deccan volcanism in relation to the Cretaceous-Paleogene boundary. Science 363 : 866– 70 [Google Scholar]
• Stanley SM. 2010 . Relation of Phanerozoic stable isotope excursions to climate, bacterial metabolism, and major extinctions. PNAS 107 : 19185– 89 [Google Scholar]
• Steffen W , Rockström J , Richardson K , Lenton TM , Folke C et al. 2018 . Trajectories of the Earth system in the Anthropocene. PNAS 115 : 8252– 59 [Google Scholar]
• Strogatz S. 1994 . Nonlinear Dynamics and Chaos New York: Addison-Wesley [Google Scholar]
• Sundquist ET 1985 . Geological perspectives on carbon dioxide and the carbon cycle. The Carbon Cycle and Atmospheric CO 2 : Natural Variations Archean to Present ET Sundquist, WS Broecker 55– 59 Washington, DC: Am. Geophys. Union [Google Scholar]
• Sundquist ET. 1990 . Influence of deep-sea benthic processes on atmospheric CO 2 . Philos. Trans. R. Soc. Lond. A 331 : 155– 65 [Google Scholar]
• Sundquist ET. 1991 . Steady-and non-steady-state carbonate-silicate controls on atmospheric CO 2 . Quat. Sci. Rev. 10 : 283– 96 [Google Scholar]
• Tyndall J. 1863 . On radiation through the earth's atmosphere. Lond. Edinb. Dublin Philos. Mag. J. Sci. 25 : 200– 6 [Google Scholar]
• Tyrrell T. 2020 . Chance played a role in determining whether Earth stayed habitable. Nat. Commun. Earth Environ. 1 : 61 [Google Scholar]
• Tziperman E , Raymo ME , Huybers P , Wunsch C. 2006 . Consequences of pacing the Pleistocene 100 kyr ice ages by nonlinear phase locking to Milankovitch forcing. Paleoceanography 21 : PA4206 [Google Scholar]
• Volterra V. 1926 . Variazioni e fluttuazioni del numero d'individui in specie animali conviventi. Mem. R. Accad. Lincei 2 : 31– 113 [Google Scholar]
• Walker JC , Hays P , Kasting JF. 1981 . A negative feedback mechanism for the long-term stabilization of Earth's surface temperature. J. Geophys. Res. Oceans 86 : 9776– 82 [Google Scholar]
• Walliser OH 1996 . Global Events and Event Stratigraphy in the Phanerozoic Berlin: Springer [Google Scholar]
• Wallmann K , Flögel S , Scholz F , Dale AW , Kemena TP et al. 2019 . Periodic changes in the Cretaceous ocean and climate caused by marine redox see-saw. Nat. Geosci. 12 : 456– 61 [Google Scholar]
• Westerhold T , Marwan N , Drury AJ , Liebrand D , Agnini C et al. 2020 . An astronomically dated record of Earth's climate and its predictability over the last 66 million years. Science 369 : 1383– 87 [Google Scholar]
• Westerhold T , Röhl U , Frederichs T , Agnini C , Raffi I et al. 2017 . Astronomical calibration of the Ypresian timescale: implications for seafloor spreading rates and the chaotic behavior of the solar system?. Clim. Past 13 : 1129– 52 [Google Scholar]
• Wieczorek S , Ashwin P , Luke CM , Cox PM. 2010 . Excitability in ramped systems: the compost-bomb instability. Proc. R. Soc. A 467 : 1243– 69 [Google Scholar]
• Wignall PB , Twitchett RJ. 1996 . Oceanic anoxia and the End Permian mass extinction. Science 272 : 1155– 58 [Google Scholar]
• Williams RG , Follows MJ. 2011 . Ocean Dynamics and the Carbon Cycle: Principles and Mechanisms Cambridge, UK: Cambridge Univ. Press [Google Scholar]
• Wunsch C. 2004 . Quantitative estimate of the Milankovitch-forced contribution to observed Quaternary climate change. Quat. Sci. Rev. 23 : 1001– 12 [Google Scholar]
• Zachos JC , Dickens GR , Zeebe RE. 2008 . An early Cenozoic perspective on greenhouse warming and carbon-cycle dynamics. Nature 451 : 279– 83 [Google Scholar]
• Zachos JC , Pagani M , Sloan L , Thomas E , Billups K 2001 . Trends, rhythms, and aberrations in global climate 65 Ma to present. Science 292 : 686– 93 [Google Scholar]
• Zeebe RE , Caldeira K. 2008 . Close mass balance of long-term carbon fluxes from ice-core CO 2 and ocean chemistry records. Nat. Geosci. 1 : 312– 15 [Google Scholar]
• Zeebe RE , Ridgwell A , Zachos JC. 2016 . Anthropogenic carbon release rate unprecedented during the past 66 million years. Nat. Geosci. 9 : 325– 29 [Google Scholar]
• Zeebe RE , Westbroek P. 2003 . A simple model for the CaCO 3 saturation state of the ocean: the “Strangelove,” the “Neritan,” and the “Cretan” Ocean. Geochem. Geophys. Geosyst. 4 : 1104 [Google Scholar]

Data & Media loading...

• Article Type: Review Article

Most Read This Month

Most cited most cited rss feed, ocean acidification: the other co 2 problem, climate change impacts on marine ecosystems, larval dispersal and marine population connectivity, ocean deoxygenation in a warming world, carbon cycling and storage in mangrove forests, progress in understanding harmful algal blooms: paradigm shifts and new technologies for research, monitoring, and management, sea surface temperature variability: patterns and mechanisms, centuries of human-driven change in salt marsh ecosystems, the effect of submarine groundwater discharge on the ocean, plastics in the marine environment.

Must There Be a Balance of Nature?

• Published: September 2001
• Volume 16 , pages 481–506, ( 2001 )

Cite this article

• Gregory Cooper 1  

801 Accesses

50 Citations

Explore all metrics

The balance of nature concept is an old idea that manifests itself in anumber of forms in population and community ecology. This paper focuseson population ecology, where controversy surrounding the balance ofnature takes the form of perennial debates over the significance ofdensity dependence, population regulation, and species interactions suchas competition. One of the most striking features of these debates, overthe course of the previous century in ecology, is the tendency to arguethe case on largely conceptual grounds. This paper explores twoquestions. Why this tendency to settle on conceptual grounds what is soobviously an empirical issue? Are there any good conceptual arguments tobe had in this area?

This is a preview of subscription content, log in via an institution to check access.

Access this article

Subscribe and save.

• Get 10 units per month
• Download Article/Chapter or eBook
• 1 Unit = 1 Article or 1 Chapter
• Cancel anytime

Price includes VAT (Russian Federation)

Instant access to the full article PDF.

Rent this article via DeepDyve

Institutional subscriptions

Similar content being viewed by others

How the Modern Synthesis Came to Ecology

Defining the niche for niche construction: evolutionary and ecological niches

Combining mechanism and drift in community ecology: a novel statistical mechanics approach.

Andrewartha, H.G.: 1957, ‘The Use of Conceptual Models in Population Ecology', Cold Spring Harbor Symposium on Quantitative Biology 22 , 219-232.

Google Scholar  

Andrewartha, H.G. andBirch, L.C.: 1954, The Distribution and Abundance of Animals , University of Chicago Press, Chicago.

Andrewartha, H.G. andBirch, L.C.: 1984, The Ecological Web , University of Chicago Press, Chicago.

Anonymous: 1944, ‘British Ecological Society Symposium on “The Ecology of Closely Allied Species”’, Journal of Animal Ecology 13 , 176-178.

Beatty, J.: 1995, ‘The Evolutionary Contingency Thesis', in G. Wolters andJ.G. Lennox (eds), Concepts, Theories, and Rationality in the Biological Sciences , University of Pittsburgh Press, Pittsburgh, pp. 45-81.

Berryman, A.A.: 1987, ‘Equilibrium or Nonequilibrium: Is That the Question?', Bulletin of the Ecological Society of America 68 , 500-502.

Berryman, A.A.: 1991, ‘Stabilization or Regulation:What it All Means!’, Oecologia 86 , 140-143.

Botkin, D.B.: 1990, Discordant Harmonies: A New Ecology for the Twenty-First Century , Oxford University Press, New York.

Brandon, R. andRausher, M.D.: 1996, ‘Testing Adaptationism: A Comment on Orzack and Sober', American Naturalist 148 , 189-201.

Brown, J.H.: 1995, Macroecology , University of Chicago Press, Chicago.

Cappuccino, N.: 1995, ‘Novel Approaches to the Study of Population Dynamics', in N. Cappuccino andP.W. Price (eds), Population Dynamics: New Approaches and Synthesis , Academic Press, San Diego, pp. 3-18.

Cappuccino, N. andPrice, P.W. (eds): 1995, Population Dynamics: New Approaches and Synthesis , Academic Press, San Diego.

Caswell, H.: 1978, ‘Predator-mediated Coexistence: A Nonequilibrium Model', American Naturalist 112 , 127-154.

Caswell, H.: 1988, ‘Theory and Models in Ecology: A Different Perspective', Bulletin of the Ecological Society of America 69 , 102-109.

Chesson, P.L.: 1981, ‘Models for Spatially Distributed Populations: The Effect of Withinpatch Variability', Theoretical Population Biology 19 , 288-325.

Chesson, P.L.: 1982, ‘The Stabilizing Effect of a Random Environment', Journal of Mathematical Biology 15 , 1-36.

Chesson, P.L. andCase, T.J.: 1986, ‘Overview: Nonequilibrium Community Theories: Chance, Variability, History, and Coexistence', in J. Diamond andT. Case (eds), pp. 229-239.

Chitty, D.: 1996, Do Lemmings Commit Suicide? Beautiful Hypotheses and Ugly Facts , Oxford University Press, New York.

Cold Spring Harbor Symposia on Quantitative Biology: 1957, Population Studies: Animal Ecology and Demography , No. 22, The Biological Laboratory, Cold Spring Harbor, NY.

Colwell, R.K.: 1984, ‘What& New? Community Ecology Discovers Biology', in P.W. Price,C.N. Slobodchikoff andW.S. Gaud (eds), A New Ecology: Novel Approaches to Interactive Systems , John Wiley & Sons, New York, pp. 387-396.

Connell, J.H.: 1990, “Apparent Versus “Real” Competition in Plants', in J.B. Grace andD. Tilman (eds), Perspectives on Plant Competition , Academic Press, San Diego, pp. 9-26.

Connell, J.H. andSousa, W.P.: 1983, ‘On the Evidence Needed to Judge Ecological Stability or Persistence', The American Naturalist 121 , 789-824.

Connor, E.F. andSimberloff, D.: 1986, ‘Competition, Scientific Method, and Null Models in Ecology', American Scientist 74 , 155-162.

Cooper, G.: 1990, ‘The Explanatory Tools of Theoretical Population Biology', in A. Fine,M. Forbes andL. Wessels (eds), PSA 1990 , Vol. 1, pp. 165-178.

Cooper, G.: 1993, ‘The Competition Controversy in Community Ecology', Biology and Philosophy 8 , 359-384.

Cooper, G.: 1997, ‘Theoretical Modeling and Biological Laws', in L. Darden (ed.), PSA 1996 , Supplement to Philosophy of Science , 63 : 28-35.

Cooper, G.: 1998, ‘Generalizations in Ecology: A Philosophical Taxonomy', Biology and Philosophy 13 , 555-586.

Cooper, G.: forthcoming, The Science of the Struggle for Existence: On the Foundations of Ecology , Cambridge University Press, Cambridge.

Costanza, R. andMaxwell, T.: 1994, ‘Resolution and Predictability: An Approach to the Scaling Problem', Landscape Ecology 9 , 47-57.

DeAngelis, D. L. andWaterhouse, J.C.: 1987, ‘Equilibrium and Nonequilibrium Concepts in Ecological Models', Ecological Monographs 57 , 1-21.

den Boer, P.J.: 1968, ‘Spreading the Risk and Stabilization of Animal Numbers’, Acta Biotheoretica 18 , 165-194.

den Boer, P.J.: 1986, ‘The Present Status of the Competitive Exclusion Principle', Trends in Evolutionary Ecology 1 , 25-28.

den Boer, P.J. andReddingius, J.: 1996, Regulation and Stabilitzation Paradigms in Population Ecology , Chapman and Hall, London.

Dennis, B. andTaper, B.: 1994, ‘Density Dependence in Time Series Observation of Natural Populations: Estimation and Testing', Ecological Monographs 64 , 205-224.

Diamond, J. andCase, T. (eds): 1986, Community Ecology , Harper and Row, New York.

Dobzhansky, T.: 1957, ‘Discussion', from H.G. Andrewartha, ‘The Use of Conceptual Models in Population Ecology', Cold Spring Harbor Symposia on Quantitative Biology , p. 235.

Egerton, F.N.: 1973, ‘Changing Concepts of the Balance of Nature', The Quarterly Review of Biology 48 , 322-350.

Elton, C.: 1933, The Ecology of Animals , Methuen, London.

Gilbert, F.S.: 1980, ‘The Equilibrium Theory of Island Geography: Fact or Fiction', Journal of Biogeography 7 , 209-235.

Hanski, I.: 1990, ‘Density Dependence, Regulation, and Variability in Animal Populations', in M.P. Hassell andR.M. May (eds), Population Regulation and Dynamics , The Royal Society, London, pp. 141-150.

Hanski, I.: 1991a, ‘Single-species Metapopulation Dynamics: Concepts, Models and Observations’, in M. Gilpin andI. Hanski (eds), Metapopulation Dynamics: Empirical and Theoretical Investigations , Academic Press, London, pp. 17-38.

Hanski, I.: 1991b, ‘Metapopulation Dynamics: Brief History and Conceptual Domain', Biological Journal of the Linnean Society 42 , 3-16.

Hanski, I.: 1996, ‘Metapopulation Ecology', in O.E. Rhodes, Jr.,R.K. Chesser andM.H. Smith (eds), Population Dynamics in Ecological Space and Time , University of Chicago Press, Chicago, pp. 13-43.

Hanski, I., I. Woiwod andJ. Perry: 1993, ‘Density Dependence, Population Persistence, and Largely Futile Arguments', Oecologia 95 , 595-598.

Hanski, I. andGilpin, M.E.: 1997, Metapopulation Biology: Ecology, Genetics, and Evolution , Academic Press, San Diego.

Harrison, S.: 1991, ‘Local Extinction in a Metapopulation Context: An Empirical Evaluation', Biological Journal of the Linnean Society 42 , 73-88.

Holyoak, M. andLawton, J.H.: 1993, ‘Comments Arising from a Paper by Wolda and Dennis: Using and Interpreting the Results of Tests for Density Dependence', Oecologia 95 , 592-594.

Howard, L.O. andFiske, W.F.: 1911, The Importation into the United States of the Parasites of the Gipsy Moth and the Brown-Tail Moth . U.S. Department of Agriculture, Bureau of Entomology, Bulletin No. 91. Reprinted: 1977), Arno Press, New York.

Hunter, M.D. andPrice, P.W.: 1998, ‘Cycles in Insect Populations: Delayed Density Dependence or Exogenous Driving Variables?’ Ecological Entomology 23 , 216-222.

Hutchinson, G.E.: 1948, ‘Circular Causal Systems in Ecology', Annals of the New York Academy of Sciences 50 , 221-246.

Kingsland, S.: 1985, Modeling Nature . University of Chicago Press, Chicago.

Kingsland, S.: 1995, Modeling Nature , 2nd edn. University of Chicago Press, Chicago.

Kuhn, T.: 1962, The Structure of Scientific Revolutions , University of Chicago Press, Chicago.

Lack, D.L.: 1954, The Natural Regulation of Animal Numbers , Clarendon Press, Oxford.

Lakatos, I.: 1970, ‘Falsification and the Methodology of Scientific Research Programmes', in I. Lakatos andA. Musgrave (eds), Criticism and the Growth of Knowledge , Cambridge University Press, London, pp. 91-197.

May, R.M.: 1973, Stability and Complexity in Model Ecosystems , Princeton University Press, Princeton.

May, R.M.: 1986, ‘The Search for Patterns in the Balance of Nature: Advances and Retreats', Ecology 67 , 1115-1126.

McIntosh, R.P.: 1987, ‘Pluralism in Ecology', Annual Review of Ecological Systems 18 , 321-341.

Menge, B.A.: 1992, ‘Community Regulation: UnderWhat Conditions Are Bottom-Up Factors Important on Rocky Shores?’ Ecology 73 , 755-765.

Murdoch, W.W.: 1994, ‘Population Regulation in Theory and Practice', Ecology 75 , 271-287.

Murray, B.G.: 1982, ‘On the Meaning of Density Dependence', Oecologia 53 , 370-373.

Murray, B.G.: 1987, ‘The Calculation of the Rate of Increase, R, of Populations with Heterogeneous Life Histories', Oikos 50 , 262-266.

Murray, B.G.: 1998, ‘Can the Population Regulation Controversy Be Buried and Forgotten?’ Oikos 84 , 148-152.

Nicholson, A.J.: 1933, ‘The Balance of Animal Populations', Journal of Animal Ecology 2 , 132-178.

Nicholson, A.J.: 1954, ‘An Outline of the Dynamics of Animal Populations', Australian Journal of Zoology 2 , 9-65.

Nicholson, A.J.: 1957, ‘Discussion', from L.C. Birch, ‘The Role of Weather in Determining the Distribution and Abundance of Animals', Cold Spring Harbor Symposia on Quantitative Biology 22 , 216.

Nicholson, A.J.: 1960, ‘The Role of Population Dynamics in Natural Selection', in S. Tax (ed.), Evolution After Darwin , University of Chicago Press, Chicago.

Nicholson, A.J. andBailey, V.A.: 1935, The Balance of Animal Populations , Zoological Society of London, London.

Orians, G.H.: 1962, ‘Natural Selection and Ecological Theory', The American Naturalist 46 , 257-263.

Orzack, S.H. andSober, E.: 1994, ‘Optimality Models and the Test of Adaptationism', American Naturalist 143 , 361-380.

Pickett, S.T.A.,Kolasa, J. andJones, C.G.: 1994, Ecological Understanding , Academic Press, New York.

Price, P.W.: 1984, ‘Alternative Paradigms in Community Ecology', in P.W. Price,C.N. Slobodchikoff andW.S. Gaud (eds), pp. 354-383.

Price, P.W.,Slobodchikoff, C.N. andGaud, W.S.: 1984, A New Ecology: Novel Approaches to Interactive Systems , John Wiley & Sons, New York.

Price, P.W. andHunter, M.D.: 1995, ‘Novelty and Synthesis in the Development of Population Dynamics', in N. Cappuccino andP.W. Price (eds), pp. 389-413.

Reddingius, J.: 1971, ‘Gambling for Existence: A Discussion of Some Theoretical Problems in Animal Populations Ecology', Acta Biotheoretica 20 (suppl.), 1-208.

Reeve, J. D.,Ayres, M.P. andLorio, P.L. Jr.: 1995, ‘Host Sustainability, Predation, and Bark Beetle Population Dynamics', in N. Cappuccino andP.W. Price (eds), pp. 339-358.

Resetarits, W.J., Jr andBernardo, J.: 1998, Experimental Ecology: Issues and Perspectives , Oxford University Press, New York.

Richardson, J.L.: 1980, ‘The Organismic Community: Resilience of an Embattled Ecological Concept', BioScience 30 , 465-471.

Ricklefs, R.E.: 2000, ‘Density Dependence, Evolutionary Optimization, and the Diversification of Avian Life Histories', The Condor 102 , 9-22.

Roughgarden, J.S.D.,May, R.M. andLevin, S.A. (eds): 1989, Perspectives in Ecological Theory , Princeton University Press, Princeton.

Royama, T.: 1992, Analytical Population Dynamics , Chapman & Hall, London.

Shipley, B. andKeddy, P.A.: 1987, ‘The Individualistic and Community-Unit Concepts as Falsifiable Hypotheses', Vegetatio 69 , 47-55.

Shipley, B. andPeters, R.H.: 1991, ‘The Seduction by Mechanism: A Reply to Tilman', The American Naturalist 138 , 1276-1282.

Shrader-Frechette, K.S.: 1989, ‘Idealized Laws, Antirealism, and Applied Science: A Case in Hydrogeology', Synthese 81 , 329-352.

Shrader-Frechette, K.S. andMcCoy, E.D.: 1993, Method in Ecology , Cambridge University Press, Cambridge.

Strong, D.R.: 1984a, ‘Density-Vague Ecology and Liberal Population Regulation in Insects', in P.W. Price,C.N. Slobodchikoff and W.S. Gaud (eds), A New Ecology: Novel Approaches to Interactive Systems , John Wiley & Sons, New York, pp. 313-327.

Strong, D.R.: 1984b, ‘Natural Variability and the Manifold Mechanisms of Ecological Communities', in G.W. Salt (ed.), Ecology and Evolutionary Biology: A Round Table on Research , University of Chicago Press, Chicago, pp. 56-80.

Strong, D.R.: 1986, ‘Density Vagueness: Abiding the Variance in the Demography of Real Populations', in J. Diamond andT. Case (eds), Community Ecology , Harper and Row, New York, pp. 257-268.

Thompson, W.R.: 1939, ‘Biological Control and the Theories of the Interaction of Populations', Parasitology 31 , 299-388.

Tilman, D.: 1989, ‘Discussion: Population Dynamics and Species Interactions', in J.S.D. Roughgarden,R.M. May andS.A. Levin (eds), Perspectives in Ecological Theory , Princeton University Press, Princeton, pp. 89-100.

Tilman, D.: 1991, ‘The Schism between Theory and Ardent Empiricism: A Reply to Shipley and Peters', The American Naturalist 138 , 1283-1286.

Tilman, D.: 1993, ‘Community Diversity and Succession: The Roles of Competition, Dispersal, and Habitat Modification', in E.D. Schulze andH.A. Mooney (eds), Biodiversity and Ecosystem Function , Springer-Verlag, Berlin, pp. 328-344.

Tilman, D. andKareiva, P.: 1997, Spatial Ecology: The Role of Space in Population Dynamics and Interspecific Interactions , Monographs in Population Biology, No. 30, Princeton University Press, Princeton.

Turchin, P.: 1995, ‘Population Regulation: Old Arguments and a New Synthesis', in N. Cappuccino andP.W. Price (eds), Population Dynamics: New Approaches and Synthesis , Academic Press, San Diego, pp. 19-41.

Turchin, P.: 1999, ‘Population Regulation: A Synthetic View', Oikos 84 , 153-159.

Turchin, P.,Getz, W.M.,Taneyhill, D.E. andGinzburg, L.: 1993, ‘The Logistic Equation Revisited: Final Installment,’ in ‘Letters to the Editor', Trends in Environmental Ecology 8 , 68-70.

Turchin, P. andBerryman, A.A.: 2000, ‘Detecting Cycles and Delayed Density Dependence: A Comment on Hunter and Price (1998)', Ecological Entomology 25 , 119-121.

Walter, G.H. andPatterson, H.E.H.: 1995, ‘Levels of Understanding in Ecology: Interspecific Competition and Community Ecology', Australian Journal of Ecology 20 , 463-466.

White, T.C.R.: 1993, The Inadequate Environment: Nitrogen and the Abundance of Animals , Springer Verlag, Berlin.

Williams, G.C.: 1992, ‘“Gaia,” Nature Worship and Biocentric Fallacies', The Quarterly Review of Biology 67 , 479-486.

Wilson, D.S.: 1989, ‘Reviving the Superorganism', Journal of Theoretical Biology 136 , 337-356.

Wilson, J.: 1999, 'To Be Continued ... ', Biology and Philosophy 14 , 615-619.

Wimsatt, W.C.: 1980, ‘Randomness and Perceived-Randomness in Evolutionary Theory', in E. Saarinen (ed.), Conceptual Issues in Ecology , D. Reidel Publishing Company, Dordrecht, The Netherlands, pp. 279-322.

Wolda, H.: 1989, ‘The Equilibrium Concept and Density Dependence Tests:What Does It All Mean?’ Oecologia 81 , 430-432.

Wolda, H.: 1991, ‘The Usefulness of the Equilibrium Concept in Population Dynamics', Oecologia 86 , 144-145.

Wolda, H.,Dennis, B. andTaper, M.L.: 1994, Density Dependent Tests, and Largely Futile Comments: Answers to Holyoak and Lawton (1993) and Hanski, Woiwod and Perry (1993)', Oecologia 98 , 229-234.

Download references

Author information

Authors and affiliations.

Washington and Lee University, Lexington, VA, 24450-0303, USA

Gregory Cooper

You can also search for this author in PubMed   Google Scholar

Rights and permissions

Reprints and permissions

About this article

Cooper, G. Must There Be a Balance of Nature?. Biology & Philosophy 16 , 481–506 (2001). https://doi.org/10.1023/A:1011935220219

Download citation

Issue Date : September 2001

DOI : https://doi.org/10.1023/A:1011935220219

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

• a priori argument
• abiotic factors
• balance of nature
• biotic factors
• competition
• density dependence
• equilibrium
• non-equilibrium
• population regulation
• random walk
• statistical return tendency
• Find a journal
• Publish with us
• Track your research

• Rewilding Ethics >>

The Balance of Nature?

Snake River Wild & Scenic River, Hells Canyon Wilderness Area, Hells Canyon NRA, ID-OR (c) Dave Foreman

“This essay is dedicated to the memory of Smilodon Dave Foreman. He was a hero and I am glad I got to meet him and know him to the extent that I did. His bold powerful vision will have cascading effects for a long time to come. I feel lucky to be a small part of it all.” —Kirk Robinson

Recently I participated in an email discussion on “The Balance of Nature.” We have all heard the phrase and most of us take for granted that there is a balance of nature. But is there? Some of the discussion participants thought there is and others thought there is not. I argued that there is, though it is not easy to say what it is. I gave the example of a tightrope walker skillfully walking a tightrope for conveying the idea of a balance of nature. She must continually make minute adjustments to maintain her balance. If she loses her balance, either because of a misstep or a gust of wind, she crosses a tipping point and falls. As a result of the discussion, I read two books on the subject that were recommended to me or referenced by other participants, The Balance of Nature: Ecology’s Enduring Myth (2009) by John Kricher and Restoring the Balance (2021) by John Vucetich. Kricher argues that there is no such thing as the balance of nature; Vucetich tries to persuade us that there is.

This is an important issue for conservationists because the word ‘balance’ is value valenced: it is good to be in balance and bad to be out of balance. Therefore, so many conservationists believe, we have an ethical responsibility not to upset the balance of nature and to restore balance where we can—say, by reintroducing an extirpated apex carnivore.

It is easy to think that there must either be a balance of nature or not—that the word ‘balance’ refers to a definite state or condition that something is either in or not in. But ‘balance’ is used in various ways to speak, for example, of a bank balance, a balanced rock, a tightrope walker, and the palate of a fine tequila to mention just a few. These many different uses form a family through resemblance and need not share a common feature. An established practice of using the term, along with accepted criteria for its application, is unique to each context of use; and a speaker must master this use in order to be said to know the meaning of the term in that context. Despite this, we humans, with our craving for generality, tend to naturally presume that the word has a perfectly definite univocal meaning that it carries with it into all contexts since, after all, it is the same word that is used in all those different contexts. However, this is a false picture of linguistic meaning. The meaning of a word is partly determined by the context of its use. Therefore, if ‘balance’ is going to have meaning with respect to nature at large, that meaning must first be explained. (Imagine coming across a traffic stop sign deep in a trackless wilderness to get the point.)

This way of looking at linguistic meaning comes from the philosopher Ludwig Wittgenstein, who is widely regarded as the most important philosopher of the 20th century. It has revolutionized many academic disciplines, e.g., linguistics, psychology, sociology, history, and communications by dislodging the quaint idea that linguistic meanings are some sort of mysterious mental entities that speakers learn to associate with words and that prescribe and guide us in their correct use like a set of rules.

Because the concept of the balance of nature is not a scientific one and there is no general theory of what constitutes a balance of nature, it makes no sense to say that there is a balance of nature or that there is not unless a particular meaning is clearly specified. Lacking this, even if this or that hypothesis about how nature works does not hold up, nothing whatever follows regarding whether there is a balance of nature. For example, if Frederick Clements’s theory of the succession of plant communities is not universally true, as Kricher reports, nothing follows one way or another regarding balance of nature.

What we can say as scientists and non-scientists alike is that nature is not totally chaotic—not a “blooming, buzzing confusion,” in William James’s memorable phrase. Change occurs at all scales, but when it comes to biological systems, such as ecosystems and organisms, the change is systemic. Ecosystems are cybernetic systems. They display recurrent patterns involving feedback loops and are stable through extended periods of time. Similarly, individual animals and plants are homeostatic. And, as the Gaia hypothesis has it, Earth is a superorganism because life processes create the very conditions that support life. Such observations give rise to the idea of a balance of nature but they do not amount to a well-defined concept or scientific theory. By ‘idea’ I mean a notion that is too vague to count as an actual concept due to the lack of an established practice of use involving accepted criteria for the application of a word.

Kricher thinks the idea of the balance of nature is that of teleology à la Aristotle, meaning that an ecosystem, indeed the entire biosphere, is inexorably moving toward a final permanent state (like a ball bearing coming to rest at the bottom of a bowl) where it will remain forever unless an exogenous force intervenes – an asteroid impact, for example. Unfortunately, he never makes it clear that this is what Aristotle means by teleology. In any case, this strikes me as an unrealistic requirement for balance in nature. It does not surprise me that Kricher could find no clear instances of it. I think he commits the “straw man” fallacy.

It is not necessary to construe the balance of nature as having anything to do with teleology, and I doubt if many ecologists have done so. It is widely accepted that change originates within the biosphere itself from random gene mutations, genetic drift, and other endogenous causes, not only from exogenous causes. This change effects change in the non-living environment, which in turn effects change in the biosphere, and so on.

Conservation Ethics

Polar bear and cubs © 2017 Dave Foreman

Given that he does not believe in the balance of nature, it is no surprise that Kricher sees no reason to engage in protection or conservation of biological nature other than to protect its potential usefulness to humans. This goes even for other animals. His environmental ethic is thoroughly anthropocentric: nature derives its value from being a potential human resource. From this perspective, the only argument for conservation is prudential: Use nature prudently so that you do not impair its ability to sustain continued use! (Given this view, I wonder why a person should not regard other people strictly as resources too . . . if he thinks he can get away with it!)

This is an unsatisfactory conservation ethic. It is inadequate for grounding an ethical obligation to conserve polar bears, for example. What if the ruling powers conclude, based on facts and plausible assumptions, that doing what it takes to conserve polar bears (or any other species) will not benefit humanity (or members of the ruling powers)? Furthermore, it is highly doubtful that many people will be motivated to conserve polar bears just because they believe it might turn out to be good for future humans.

Unlike Kricher, Vucetich thinks there is something we can reasonably call the balance of nature, specifically the predator-prey relationship and the phenomenon of the trophic cascade: predators control the number and demographics of prey animals, which in turn keeps the plant community healthy and capable of supporting a diverse biota. It is well known that the slaughter of predators on the Kaibab Plateau in the early 20 th century was intended to increase the mule deer herd but had a deteriorating effect on the habitat, leading instead to mass deer starvation. That was an example of what happens when you upset the “balance” by shutting down a trophic cascade. It was also one of the reasons for reintroducing wolves to Yellowstone National Park (to control the elk population), and more recently to genetically rescue the dwindling wolf population in Isle Royale National Park (to control the moose population). Whether we want to call this restoring the balance, as Vucetich does, is not terribly important, but it has nothing to do with teleology. What is important is that an ecosystem is a self-renewing homeostatic system, analogous in this respect to an organism. In addition, Vucetich points out that all organisms are related as members of the family of life that descended from a common ancestor. In this way, he hopes to persuade readers that they have an ethical obligation to care for other creatures and their habitats just as they do for members of their own immediate family. Personally, I doubt if this alone is motivationally powerful enough to do the job.

Motivation aside, do these facts place us under an ethical obligation to care for other creatures? Am I my brother’s keeper, so to speak, even where my “brother” is a species of snail on an island in the ocean? Again, I am dubious, though this concern is largely ameliorated if my obligation is limited to matters that I can influence. Bottom line: I think this view has a lot more going for it than Kricher’s but still falls short of what we need.

However, Vucetich develops his view further by recognizing that many other creatures besides humans have interests. For example, squirrels have an interest in storing nuts; wolverines have an interest in scavenging carrion; etc. Whether they reason about what to do is beside the point. It is enough that they consciously and purposely pursue what helps protect and sustain their lives and the lives of their offspring, as well as what gives them pleasure, such as play. And just as we recognize an ethical obligation to other humans not to interfere with their ability to freely pursue their interests (with some exceptions), Vucetich thinks we ought to recognize a similar obligation with respect to other creatures.

This seems plausible enough with respect to mammals and birds, and perhaps reptiles and amphibians, but do insects, plants, and ecosystems have interests? We can say that some things are in their interest , in the sense of contributing to their survival, maintenance, or flourishing, but this is different from them literally having interests. The difference is illustrated by a bear who is interested in what’s on offer at a bait station where a “hunter” waits to kill her. What she is interested in is not in her interest. I think Vucetich tends to conflate the two senses of “having an interest,” but for the most part they no doubt tend to naturally coincide: wild animals are mostly interested in what is in their interest.

Also, while I am sympathetic to the needs of other creatures, it is not obvious to me why I am obligated to care about them enough to help protect them. After all, I have my own interests to look out for. Therefore, if I am obligated to care about the interests of other creatures, I think the reason must be more basic than the mere fact that they have interests (in either sense).

A Good Alternative 

Sandhill Cranes silhuetted in fog, Bosque del Apache NWR, NM, (c) Dave Foreman

Suppose we focus more broadly on what is good for a plant, animal, or ecosystem. It is good for a tree to receive sunshine and moisture (even though, like an ecosystem, it is presumably incapable of conscious experience or agency). It is good for a mountain lion to have cover for stalking prey. It is good for the Great Salt Lake ecosystem with its millions of migratory birds to receive inflow from rivers and streams. This use of ‘good’ is different from saying, for example, that it is good for a knife to be sharp. It makes no difference to a knife whether it is sharp, only to the user of the knife. We might express this by saying that plants, animals, and ecosystems have a good of their own. A knife does not have a good of its own. It’s just a thing.

Following Aristotle, it makes sense to say that what is good for a creature is what is in accord with its nature. It is good for a golden retriever to have a compassionate human caretaker. It is good for a wolf to live in the wild (unless it has been raised in captivity), even though it might starve or get a broken jaw from the kick of an elk. And similarly for an elk, even though it might get killed by wolves. Life in the wild is the best possible life for both wolf and elk because that is what evolution has equipped them for and what allows them maximum potential to flourish as the kinds of creature they are. And this is itself good—not instrumentally good as in “good for,” but good in the sense of being fitting or right.

Now to the crucial question: Do we humans have an ethical obligation to not damage what is good? Clearly, a justification is needed—and a good one—for purposely doing so. I do not see how this can be cogently denied. The sentence “It is not wrong to wantonly damage what is good” is close to an outright contradiction. Furthermore, if we humans have an obligation to not damage what is good without good reason, then, I submit, we also have an obligation to repair what we have damaged where we can. I think this is axiomatic. If not, then what is the point of the obligation to not damage in the first place? (Vucetich argues to the same conclusion in a more roundabout way based on a theory of restorative justice.)

This view is consistent with Vucetich’s point about interests but goes further by (1) more explicitly including plant communities such as old growth forests, and ecosystems, in the class of things to which we have an ethical obligation; (2) by grounding the obligation more firmly and transparently in what is good; and (3) by directly implying an obligation to undertake rewilding efforts where we can, even when we are faced with a competing obligation to not harm individual creatures—as when it must be decided whether or not to lethally remove an invasive species that is causing damage to an ecosystem. In such cases, we must decide which of competing obligations ought to take precedence while keeping in mind that “goods-of-its-own” are not necessarily all equally good in an absolute sense. There is a vast difference between pulling weeds in your garden and clear-cutting an old growth forest, for example; and between swatting a housefly and murder. However, this complicated issue lies beyond the scope of this essay.

If someone is convinced, based on established facts and sound reasoning, that she has an ethical obligation to do (or not do) something, it can motivate her to act accordingly—even against a powerful contrary inclination. As well, the best available science and sound ethical reasoning should inform and guide government conservation policies & programs. But the overarching challenge for us conservationists is to foster the growth of a cultural conservation ethos à la Aldo Leopold’s “land ethic” that also teaches compassion for individual creatures. I hope this essay imparts some impetus toward that goal.

Kirk grew up in Bountiful, Utah between the shore of the Great Salt Lake and the foothills of the Wasatch Mountains. As a child, he loved roaming the foothills looking for wild animals and visiting Farmington Bay Bird Refuge. Naturally, he fell in love with the land and its wildlife. Since those early years, he has spent a big part of his life exploring the deserts, rivers, and mountains of the West.

In the 1990s, Kirk and some friends from the Utah Wilderness Association began working to reform Utah wildlife governance and management to make it more democratic, ecologically sound, and compassionate. This led to the founding of a non-profit organization, Western Wildlife Conservancy , to address the issues. Of particular concern is the scientifically and ethically misguided way that native carnivores such as mountain lions, black bears, and gray wolves are treated. The vital role that these intelligent and magnificent creatures play in maintaining the health of ecosystems goes unappreciated, as evidenced by a long history of persecution. In addition to being Executive Director of Western Wildlife Conservancy, Kirk is on the Leadership Council of The Rewilding Institute and the Advisory Committee of Wildlife For All.

Prior to Western Wildlife Conservancy, Kirk was a Professor of Philosophy. He has a PhD in Philosophy from the University of Cincinnati and has taught at universities in Montana and Utah. In 2004, he graduated from the S.J. Quinney College of Law at the University of Utah with a certificate in Natural Resource Law to better equip him for work on wildlife conservation issues. In his free time, he enjoys backpacking, x-c skiing, river trips, playing the acoustic guitar, and time spent with friends and with his dog Bingo.

Related Rewilding Earth Articles

Ethics of Wildlife Conservation, Part 3: The Good

The Ethics of Wildlife Conservation: Part 2

The Ethics of Wildlife Conservation

Insert/edit link

Enter the destination URL

Or link to existing content

Thank you for visiting nature.com. You are using a browser version with limited support for CSS. To obtain the best experience, we recommend you use a more up to date browser (or turn off compatibility mode in Internet Explorer). In the meantime, to ensure continued support, we are displaying the site without styles and JavaScript.

• View all journals
• Explore content
• About the journal
• Publish with us
• Sign up for alerts
• Published: 03 September 2024

Stage dependence of Elton’s biotic resistance hypothesis of biological invasions

• Kun Guo   ORCID: orcid.org/0000-0001-9597-2977 1 ,
• Petr Pyšek   ORCID: orcid.org/0000-0001-8500-442X 2 , 3 ,
• Milan Chytrý   ORCID: orcid.org/0000-0002-8122-3075 4 ,
• Jan Divíšek   ORCID: orcid.org/0000-0002-5127-5130 4 , 5 ,
• Martina Sychrová 4 , 5 ,
• Zdeňka Lososová 4 ,
• Mark van Kleunen   ORCID: orcid.org/0000-0002-2861-3701 6 , 7 ,
• Simon Pierce 8 &
• Wen-Yong Guo   ORCID: orcid.org/0000-0002-4737-2042 1 , 9 , 10  

Nature Plants ( 2024 ) Cite this article

Metrics details

• Biodiversity
• Invasive species

Elton’s biotic resistance hypothesis posits that species-rich communities are more resistant to invasion. However, it remains unknown how species, phylogenetic and functional richness, along with environmental and human-impact factors, collectively affect plant invasion as alien species progress along the introduction–naturalization–invasion continuum. Using data from 12,056 local plant communities of the Czech Republic, this study reveals varying effects of these factors on the presence and richness of alien species at different invasion stages, highlighting the complexity of the invasion process. Specifically, we demonstrate that although species richness and functional richness of resident communities had mostly negative effects on alien species presence and richness, the strength and sometimes also direction of these effects varied along the continuum. Our study not only underscores that evidence for or against Elton’s biotic resistance hypothesis may be stage-dependent but also suggests that other invasion hypotheses should be carefully revisited given their potential stage-dependent nature.

This is a preview of subscription content, access via your institution

Access options

Access Nature and 54 other Nature Portfolio journals

Get Nature+, our best-value online-access subscription

24,99 € / 30 days

cancel any time

Subscribe to this journal

Receive 12 digital issues and online access to articles

111,21 € per year

only 9,27 € per issue

Buy this article

• Purchase on SpringerLink
• Instant access to full article PDF

Prices may be subject to local taxes which are calculated during checkout

Similar content being viewed by others

Native diversity buffers against severity of non-native tree invasions

Invasion intensity influences scale-dependent effects of an exotic species on native plant diversity

The impact of land use on non-native species incidence and number in local assemblages worldwide

Data availability.

The data used in this study were obtained from these sources: the data on vegetation plots were from the Czech National Phytosociological Database 54 ( https://botzool.cz/vegsci/phytosociologicalDb/ ); species’ statuses along the invasion continuum were extracted from Pyšek et al. 55 ; the three leaf traits required for CSR calculation were collected from the Pladias Database of the Czech Flora and Vegetation 58 and other publications 61 , 62 , 63 , 64 , 65 , 66 ; species CSR scores were calculated using the StrateFy tool 60 ; climatic variables were extracted from Tolasz 73 ; soil pH was collected from the Land Use/Land Cover Area Frame Survey 74 ; and the human population density of the cadastral area where each plot located was obtained from the Digital Vector Database of Czech Republic ArcČR v.4.0 (ref. 75 ). The data that support the findings of this study are available via GitHub at https://github.com/kun-ecology/BioticResistance_InvasionContinuum and via Zenodo at https://doi.org/10.5281/zenodo.12818669 (ref. 79 ).

Code availability

R functions for the computation of phylogenetic and functional metrics have been deposited on GitHub ( https://github.com/kun-ecology/ecoloop ). R scripts for reproducing the analyses and figures are available via GitHub at https://github.com/kun-ecology/BioticResistance_InvasionContinuum and via Zenodo at https://doi.org/10.5281/zenodo.12818669 (ref. 79 ).

Roy, H. E. et al. (eds) Summary for Policymakers of the Thematic Assessment Report on Invasive Alien Species and Their Control of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (IPBES Secretariat, 2023).

Spatz, D. R. et al. Globally threatened vertebrates on islands with invasive species. Sci. Adv. 3 , e1603080 (2017).

Article   PubMed   PubMed Central   Google Scholar  

Walsh, J. R., Carpenter, S. R. & Vander Zanden, M. J. Invasive species triggers a massive loss of ecosystem services through a trophic cascade. Proc. Natl Acad. Sci. USA 113 , 4081–4085 (2016).

Article   CAS   PubMed   PubMed Central   Google Scholar  

Liu, C. et al. Economic costs of biological invasions in Asia. NeoBiota 67 , 53–78 (2021).

Article   Google Scholar  

Capinha, C., Essl, F., Porto, M. & Seebens, H. The worldwide networks of spread of recorded alien species. Proc. Natl Acad. Sci. USA 120 , e2201911120 (2023).

Article   CAS   PubMed   Google Scholar  

Seebens, H. et al. Projecting the continental accumulation of alien species through to 2050. Glob. Change Biol. 27 , 970–982 (2021).

Article   CAS   Google Scholar  

Richardson, D. M. & Pyšek, P. Plant invasions: merging the concepts of species invasiveness and community invasibility. Prog. Phys. Geogr. 30 , 409–431 (2006).

Elton, C. S. The Ecology of Invasions by Animals and Plants (Univ. Chicago Press, 1958).

Bach, W. et al. Phylogenetic composition of native island floras influences naturalized alien species richness. Ecography 2022 , e06227 (2022).

Beaury, E. M., Finn, J. T., Corbin, J. D., Barr, V. & Bradley, B. A. Biotic resistance to invasion is ubiquitous across ecosystems of the United States. Ecol. Lett. 23 , 476–482 (2020).

Article   PubMed   Google Scholar  

Lefebvre, S., Segar, J. & Staude, I. R. Non-natives are linked to higher plant diversity across spatial scales. J. Biogeogr. 51 , 1290–1298 (2024).

Fridley, J. D. et al. The invasion paradox: reconciling pattern and process in species invasion. Ecology 88 , 3–17 (2007).

Herben, T., Mandák, B., Bímová, K. & Münzbergová, Z. Invasibility and species richness of a community: a neutral model and a survey of published data. Ecology 85 , 3223–3233 (2004).

Jeschke, J. M. et al. Taxonomic bias and lack of cross-taxonomic studies in invasion biology. Front. Ecol. Environ. 10 , 349–350 (2012).

Jeschke, J. M. General hypotheses in invasion ecology. Divers. Distrib. 20 , 1229–1234 (2014).

Stohlgren, T. J., Barnett, D. T. & Kartesz, J. T. The rich get richer: patterns of plant invasions in the United States. Front. Ecol. Environ. 1 , 11–14 (2003).

Shea, K. & Chesson, P. Community ecology theory as a framework for biological invasions. Trends Ecol. Evol. 17 , 170–176 (2002).

Delavaux, C. S. et al. Native diversity buffers against severity of non-native tree invasions. Nature 621 , 773–781 (2023).

Blackburn, T. M. et al. A proposed unified framework for biological invasions. Trends Ecol. Evol. 26 , 333–339 (2011).

Pyšek, P. & Richardson, D. M. Invasive species, environmental change and management, and health. Annu. Rev. Environ. Resour. 35 , 25–55 (2010).

Daly, E. Z. et al. A synthesis of biological invasion hypotheses associated with the introduction–naturalisation–invasion continuum. Oikos 2023 , e09645 (2023).

Guo, K. et al. Ruderals naturalize, competitors invade: varying roles of plant adaptive strategies along the invasion continuum. Funct. Ecol. 36 , 2469–2479 (2022).

Pyšek, P. et al. Small genome size and variation in ploidy levels support the naturalization of vascular plants but constrain their invasive spread. N. Phytol. 239 , 2389–2403 (2023).

Omer, A. et al. The role of phylogenetic relatedness on alien plant success depends on the stage of invasion. Nat. Plants 8 , 906–914 (2022).

Guo, K. et al. Plant invasion and naturalization are influenced by genome size, ecology and economic use globally. Nat. Commun. 15 , 1330 (2024).

Richardson, D. M. & Pyšek, P. Naturalization of introduced plants: ecological drivers of biogeographical patterns. N. Phytol. 196 , 383–396 (2012).

Byun, C., De Blois, S. & Brisson, J. Plant functional group identity and diversity determine biotic resistance to invasion by an exotic grass. J. Ecol. 101 , 128–139 (2013).

Banerjee, A. K. et al. Not just with the natives, but phylogenetic relationship between stages of the invasion process determines invasion success of alien plant species. Preprint at https://doi.org/10.1101/2022.10.12.512006 (2022).

Cubino, J. P., Těšitel, J., Fibich, P., Lepš, J. & Chytrý, M. Alien plants tend to occur in species-poor communities. NeoBiota 73 , 39–56 (2022).

Lannes, L. S. et al. Species richness both impedes and promotes alien plant invasions in the Brazilian Cerrado. Sci. Rep. 10 , 11365 (2020).

Stohlgren, T. J. et al. Exotic plant species invade hot spots of native plant diversity. Ecol. Monogr. 69 , 25–46 (1999).

Davidson, A. M., Jennions, M. & Nicotra, A. B. Do invasive species show higher phenotypic plasticity than native species and, if so, is it adaptive? A meta-analysis. Ecol. Lett. 14 , 419–431 (2011).

Lau, J. A. & Funk, J. L. How ecological and evolutionary theory expanded the ‘ideal weed’ concept. Oecologia 203 , 251–266 (2023).

Prentis, P. J., Wilson, J. R. U., Dormontt, E. E., Richardson, D. M. & Lowe, A. J. Adaptive evolution in invasive species. Trends Plant Sci. 13 , 288–294 (2008).

Stohlgren, T., Jarnevich, C. S., Chong, G. W. & Evangelista, P. Scale and plant invasions: a theory of biotic acceptance. Preslia 78 , 405–426 (2006).

Google Scholar  

Cavieres, L. A. Facilitation and the invasibility of plant communities. J. Ecol. 109 , 2019–2028 (2021).

Catford, J. A., Vesk, P. A., Richardson, D. M. & Pyšek, P. Quantifying levels of biological invasion: towards the objective classification of invaded and invasible ecosystems. Glob. Change Biol. 18 , 44–62 (2012).

Su, G., Mertel, A., Brosse, S. & Calabrese, J. M. Species invasiveness and community invasibility of North American freshwater fish fauna revealed via trait-based analysis. Nat. Commun. 14 , 2332 (2023).

Parker, J. D. et al. Do invasive species perform better in their new ranges? Ecology 94 , 985–994 (2013).

Iseli, E. et al. Rapid upwards spread of non-native plants in mountains across continents. Nat. Ecol. Evol. 7 , 405–413 (2023).

Pauchard, A. et al. Ain’t no mountain high enough: plant invasions reaching new elevations. Front. Ecol. Environ. 7 , 479–486 (2009).

Zheng, M.-M., Pyšek, P., Guo, K., Hasigerili, H. & Guo, W.-Y. Clonal alien plants in the mountains spread upward more extensively and faster than non-clonal. Neobiota 91 , 29–48 (2024).

Zu, K. et al. Elevational shift in seed plant distributions in China’s mountains over the last 70 years. Glob. Ecol. Biogeogr. 32 , 1098–1112 (2023).

Reeve, S. et al. Rare, common, alien and native species follow different rules in an understory plant community. Ecol. Evol. 12 , e8734 (2022).

Hartemink, A. E. & Barrow, N. J. Soil pH–nutrient relationships: the diagram. Plant Soil 486 , 209–215 (2023).

Dawson, W., Rohr, R. P., van Kleunen, M. & Fischer, M. Alien plant species with a wider global distribution are better able to capitalize on increased resource availability. N. Phytol. 194 , 859–867 (2012).

Dawson, W. et al. Global hotspots and correlates of alien species richness across taxonomic groups. Nat. Ecol. Evol. 1 , 0186 (2017).

Pyšek, P. et al. Disentangling the role of environmental and human pressures on biological invasions across Europe. Proc. Natl Acad. Sci. USA 107 , 12157–12162 (2010).

Damgaard, C. A critique of the space-for-time substitution practice in community ecology. Trends Ecol. Evol. 34 , 416–421 (2019).

Hejda, M., Pyšek, P. & Jarošík, V. Impact of invasive plants on the species richness, diversity and composition of invaded communities. J. Ecol. 97 , 393–403 (2009).

Lovell, R. S. L., Collins, S., Martin, S. H., Pigot, A. L. & Phillimore, A. B. Space-for-time substitutions in climate change ecology and evolution. Biol. Rev. 98 , 2243–2270 (2023).

Thomaz, S. M. et al. Using space-for-time substitution and time sequence approaches in invasion ecology. Freshw. Biol. 57 , 2401–2410 (2012).

Wogan, G. O. U. & Wang, I. J. The value of space‐for‐time substitution for studying fine‐scale microevolutionary processes. Ecography 41 , 1456–1468 (2018).

Chytrý, M. & Rafajová, M. Czech National Phytosociological Database: basic statistics of the available vegetation-plot data. Preslia 75 , 1–15 (2003).

Pyšek, P. et al. Catalogue of alien plants of the Czech Republic (3rd edition): species richness, status, distributions, habitats, regional invasion levels, introduction pathways and impacts. Preslia 94 , 447–577 (2022).

Divíšek, J. et al. Similarity of introduced plant species to native ones facilitates naturalization, but differences enhance invasion success. Nat. Commun. 9 , 4631 (2018).

Raunkiaer, C. The Life Forms of Plants and Statistical Plant Geography (Clarendon, 1934).

Chytrý, M. et al. Pladias Database of the Czech flora and vegetation. Preslia 93 , 1–87 (2021).

Li, D. rtrees: an R package to assemble phylogenetic trees from megatrees. Ecography 2023 , e06643 (2023).

Pierce, S. et al. A global method for calculating plant CSR ecological strategies applied across biomes world-wide. Funct. Ecol. 31 , 444–457 (2017).

Díaz, S. et al. The global spectrum of plant form and function: enhanced species-level trait dataset. Sci. Data 9 , 755 (2022).

Guo, W.-Y. et al. The role of adaptive strategies in plant naturalization. Ecol. Lett. 21 , 1380–1389 (2018).

Guo, W.-Y. et al. Domestic gardens play a dominant role in selecting alien species with adaptive strategies that facilitate naturalization. Glob. Ecol. Biogeogr. 28 , 628–639 (2019).

Tavşanoǧlu, Ç. & Pausas, J. G. A functional trait database for Mediterranean Basin plants. Sci. Data 51 , 180135 (2018).

Bjorkman, A. D. et al. Tundra Trait Team: a database of plant traits spanning the tundra biome. Glob. Ecol. Biogeogr. 27 , 1402–1411 (2018).

Kattge, J. et al. TRY plant trait database—enhanced coverage and open access. Glob. Change Biol. 26 , 119–188 (2020).

Goolsby, E. W., Bruggeman, J. & Ané, C. Rphylopars: fast multivariate phylogenetic comparative methods for missing data and within-species variation. Methods Ecol. Evol. 8 , 22–27 (2017).

Matthews, T. J. et al. A global analysis of avian island diversity–area relationships in the Anthropocene. Ecol. Lett. 26 , 965–982 (2023).

Cardoso, P. et al. Calculating functional diversity metrics using neighbor-joining trees. Ecography https://doi.org/10.1101/2022.11.27.518065 (2022).

Cardoso, P., Rigal, F. & Carvalho, J. C. BAT—Biodiversity Assessment Tools, an R package for the measurement and estimation of alpha and beta taxon, phylogenetic and functional diversity. Methods Ecol. Evol. 6 , 232–236 (2015).

Swenson, N. G. Functional and Phylogenetic Ecology in R (Springer, 2014).

Loiola, P. P. et al. Invaders among locals: alien species decrease phylogenetic and functional diversity while increasing dissimilarity among native community members. J. Ecol. 106 , 2230–2241 (2018).

Tolasz, R. et al. Atlas Podnebí Česka—Climate Atlas of Czechia (Czech Hydrometeorological Institute and Palacký Univ., 2007).

Ballabio, C. et al. Mapping LUCAS topsoil chemical properties at European scale using Gaussian process regression. Geoderma 355 , 113912 (2019).

Digital Vector Database of Czech Republic ArcČR v.4.0 (ČÚZK, ČSÚ and ArcDATA Prague, 2021).

Lindgren, F. & Rue, H. Bayesian spatial modelling with R-INLA. J. Stat. Softw. 63 , 1–25 (2015).

Schielzeth, H. Simple means to improve the interpretability of regression coefficients. Methods Ecol. Evol. 1 , 103–113 (2010).

R Core Team. R: A Language and Environment for Statistical Computing http://www.R-project.org/ (R Foundation for Statistical Computing, 2023).

Guo, K. et al. Stage dependence of Elton’s biotic resistance hypothesis of biological invasions. Zenodo https://doi.org/10.5281/zenodo.12818669 (2024).

Download references

Acknowledgements

K.G. and W.-Y.G. were supported by the Natural Science Foundation of China (grant no. 32171588, awarded to W.-Y.G.) and the Shanghai Pujiang Program (grant no. 21PJ1402700, awarded to W.-Y.G.). K.G. was also supported by the Shanghai Sailing Program (grant no. 22YF1411700) and the Natural Science Foundation of China (grant no. 32301386). P.P. was supported by the Czech Science Foundation (EXPRO grant no. 19-28807X) and the Czech Academy of Sciences (long-term research development project RVO 67985939). M.C. and Z.L. were supported by the Czech Science Foundation (EXPRO grant no. 19-28491X). J.D. was supported by the Technology Agency of the Czech Republic (grant no. SS02030018). M.S. was funded by the project GEOSANT with the funding organization Masaryk University (MUNI/A/1469/2023).

Author information

Authors and affiliations.

Zhejiang Tiantong Forest Ecosystem National Observation and Research Station, Shanghai Key Lab for Urban Ecological Processes and Eco-Restoration & Research Center for Global Change and Complex Ecosystems, School of Ecological and Environmental Sciences, East China Normal University, Shanghai, People’s Republic of China

Kun Guo & Wen-Yong Guo

Department of Invasion Ecology, Institute of Botany, Czech Academy of Sciences, Průhonice, Czech Republic

Department of Ecology, Faculty of Science, Charles University, Prague, Czech Republic

Department of Botany and Zoology, Faculty of Science, Masaryk University, Brno, Czech Republic

Milan Chytrý, Jan Divíšek, Martina Sychrová & Zdeňka Lososová

Department of Geography, Faculty of Science, Masaryk University, Brno, Czech Republic

Jan Divíšek & Martina Sychrová

Ecology, Department of Biology, University of Konstanz, Konstanz, Germany

Mark van Kleunen

Zhejiang Provincial Key Laboratory of Plant Evolutionary Ecology and Conservation, Taizhou University, Taizhou, People’s Republic of China

Department of Agricultural and Environmental Sciences (DiSAA), University of Milan, Milan, Italy

Simon Pierce

Zhejiang Zhoushan Island Ecosystem Observation and Research Station, Zhoushan, People’s Republic of China

Wen-Yong Guo

State Key Laboratory of Estuarine and Coastal Research, East China Normal University, Shanghai, People’s Republic of China

You can also search for this author in PubMed   Google Scholar

Contributions

K.G., P.P. and W.-Y.G. conceptualized the research. P.P., M.C., J.D., M.S. and W.-Y.G. provided the data. K.G. analysed the data and drafted the paper, with substantial contributions from all authors.

Corresponding author

Correspondence to Wen-Yong Guo .

Ethics declarations

Competing interests.

The authors declare no competing interests.

Peer review

Peer review information.

Nature Plants thanks Martha Hoopes and Johannes Kollmann for their contribution to the peer review of this work.

Additional information

Publisher’s note Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations.

Supplementary information

Supplementary information.

Supplementary Figs. 1–8.

Reporting Summary

Rights and permissions.

Springer Nature or its licensor (e.g. a society or other partner) holds exclusive rights to this article under a publishing agreement with the author(s) or other rightsholder(s); author self-archiving of the accepted manuscript version of this article is solely governed by the terms of such publishing agreement and applicable law.

Reprints and permissions

About this article

Cite this article.

Guo, K., Pyšek, P., Chytrý, M. et al. Stage dependence of Elton’s biotic resistance hypothesis of biological invasions. Nat. Plants (2024). https://doi.org/10.1038/s41477-024-01790-0

Download citation

Received : 18 April 2024

Accepted : 15 August 2024

Published : 03 September 2024

DOI : https://doi.org/10.1038/s41477-024-01790-0

Share this article

Anyone you share the following link with will be able to read this content:

Sorry, a shareable link is not currently available for this article.

Provided by the Springer Nature SharedIt content-sharing initiative

Quick links

• Explore articles by subject
• Guide to authors
• Editorial policies

Sign up for the Nature Briefing newsletter — what matters in science, free to your inbox daily.