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Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis

Published online by Cambridge University Press:  01 December 2025

Andrea Cardini*
Affiliation:
Dipartimento di Scienze Chimiche e Geologiche, Università di Modena e Reggio Emilia , Modena, Italy School of Anatomy, Physiology and Human Biology, The University of Western Australia, Crawley, WA, Australia
*
Corresponding author: Andrea Cardini; Email: andrea.cardini@uwa.edu.au
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Abstract

Scientists still debate whether small groups of Paleolithic hunter-gatherers caused the extinction of large Ice Age animals like prehistoric elephants, giant sloths and cave lions. Beyond paleontology, this question has deep sociological implications and is relevant for how we understand the role of humankind in today’s environmental crisis. A human-driven megafauna extinction has often fostered the idea of a naturalization of human environmental impacts and the belief that all people (modern or ancient, rich or poor, from any part of the world) share responsibility for the current crisis. But is that true? In the review, I discuss whether a long evolutionary history of impacts really makes us inevitably destructive, compelling humanity to accept a devastating anthropocentric dominance as the fateful destiny natural selection built for us. In contrast, I argue that, while our exceptional ability to shape environments has made us a ‘hyper-keystone’ species, benefiting only a few species and humans, this same ability also has the potential to help us restore balance to the world. That requires rejecting anthropocentric supremacy and placing ecosystems at the center stage of our relationship with nonhuman nature. We may have wiped out the mammoths and mastodons, but human destructiveness is not fate.

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Overview Review
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This is an Open Access article, distributed under the terms of the Creative Commons Attribution licence (http://creativecommons.org/licenses/by/4.0), which permits unrestricted re-use, distribution and reproduction, provided the original article is properly cited.
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© The Author(s), 2025. Published by Cambridge University Press

Impact statement

Mammoths, woolly rhinoceros, cave lions, thunder-birds, giant species of sloths, lizards and tortoises, as well as many other large terrestrial animals, including our own sister species – the Neanderthal – disappeared forever toward the end of the Ice Age. This is the largest extinction we know to specifically wipe out species of animals as large as or larger than an adult human (the so-called ‘megafauna’). The role of modern humans in this event is still debated in the sciences and humanities. Indeed, to many scholars, a human-caused extinction of the majority of the most charismatic Ice Age land vertebrates seems crucial for shaping our response to the current global environmental crisis. If true, they argue, we cannot blame modernity for our devastating impacts, as human destructiveness starts in the Stone Age and is part of what we are. Thus, a significant role of a small population of ‘low-tech’ hunter-gatherers in the pre-agricultural megafauna collapse implies the inevitability of destructive human impacts and also equal responsibility for today’s biodiversity crisis. In fact, this is a biased, deterministic view of ‘human nature’ that ignores scientific evidence and leads to the belief that modern humans can never coexist within a balanced natural ecosystem. In contrast, I maintain that human-driven global change reflects the interaction of biological and cultural evolution. Genetic change is slow, but culture is rapidly modified, and the direction of cultural change is not meant to be always negative for the ecosphere. Then, can we shift from harmful to beneficial impact by adopting a global ecosystem perspective? An objective interpretation of scientific evidence on human behavioral ecology suggests that the very abilities that make us a ‘hyper-keystone’ species are fundamental for restoring global ecosystem balance.

Introduction

Tracing human supremacy from prehistoric extinctions to modern ecological and social domination: From megafauna to Native Peoples, ‘ubi major, minor cessat’

‘One race of men — as in races of lower animals — shall disappear off the face of the earth and give place to another race … .The races of the Mammoths and Mastodons, and great Sloths, came and passed away: the Red Man of America is passing away’ (General J. H. Carleton, commanding officer in the mid-18th-century US campaign against the Apache and Navajo, quoted by Ghosh [Reference Ghosh2021, p. 51]). As the Romans had it, ‘Ubi major, minor cessat’: the weaker vanishes as someone more powerful arrives. For General Carleton, Native Americans were, thus, destined to fade away, replaced by White invaders. Thousands of years before, as the Pleistocene (or ‘Ice Age’) was coming to an end and humans colonized the Americas, the largest animals of the New World met a similar fate of decline and extinction.

Today, few would accept genocide and ecocide as an inevitability of being humans. Yet, human supremacy and the domination of ‘nature’ are embedded in the leading worldview of our time (Crist, Reference Crist2019). The presumption of human uniqueness creates an unsurpassable divide between us and the rest of nature. Not everybody shares this perspective, which, nonetheless, prevails in modern socio-political and economic systems (Ruuska et al., Reference Ruuska, Heikkurinen and Wilén2020). But what is the origin of human supremacism? Is it an existential cultural byproduct or is it an essential emergent trait of our biological evolution? In this context, can the extinction of mammoths and giant sloths be any relevant, beyond the disheartening similitude between the trajectory of population decline of both the largest animals of North America and its original inhabitants?

Perhaps, these are philosophical questions of little interest to biologists. Yet, when a student of my mammalogy class asked me where I stood in the debate that followed the first killing of a person by a brown bear in Italy, my instinctive reply was: ‘Shall we live in a world where there is just one dominant species or do we want a balanced ecosystem enriched in its functions and diversity?’. Back then, however, I had not appreciated that the demise of native populations conquered by a technologically superior invader is just a special within-species case of a broader phenomenon of human ecological dominance (Ghosh, Reference Ghosh2021). Thus, anthropogenic impacts on the environment do have potential sociological ramifications, which are also relevant for life and environmental scientists.

Carleton draws a daring (but not unusual in his time; Sayre, Reference Sayre2001) parallel between the massacre of Native Americans and the extinctions of the most charismatic species of the Ice Age, such as mammoths and other gigantic beasts, whose existence is now only known through fossil remains. In fact, alongside these giants, many other large (≥45 kg) land animals, the so-called ‘megafauna’, went extinct between 50,000 and 10,000 years ago (ya) (Koch and Barnosky, Reference Koch and Barnosky2006; Stuart, Reference Stuart2015). The primary wave of extinctions, spanning ~40,000 years, mostly coincides with the Upper Paleolithic (~50,000–12,000 ya) and, thus, the end of the Pleistocene. However, this event is usually referred to as the Late Quaternary Extinctions (LQEs), because the extinction wave extended into the early Holocene (~12,000 ya to the present). The Holocene epoch, along with the Pleistocene, forms the quaternary period (~2.6 million ya to the present), the shortest and most recent of the ‘post-dinosaur’ Cenozoic era beginning 66 million ya. It is largely because of the LQE that today’s living terrestrial megafauna is downsized and impoverished (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016; Galetti et al., Reference Galetti, Moleón, Jordano, Pires, Guimarães, Pape, Nichols, Hansen, Olesen, Munk, de Mattos, Schweiger, Owen-Smith, Johnson, Marquis and Svenning2018). Thus, in the often heated discussion on the conflict that accompanies the recovery of the European populations of brown bears and wolves (Gross, Reference Gross2023a), we seem to forget that these species are the only two survivors among all the large and diverse carnivores that once roamed this continent.

Carleton’s parallel must imply humans as the cause of the megafauna extinctions to suggest that human dominance is key to both the destruction of native populations and ecosystems. Seen from the angle of the global environmental crisis of our time, that would also indicate a continuity between prehistoric extinctions of keystone megafaunal species and later anthropogenic impacts. It is precisely this continuity that a current of sociological research finds rich in potential societal and ecological ramifications. However, these implications are often overlooked in what we, as biologists, teach about conservation and environmental change, even though they are central to the debate on how we should respond to the current crisis.

Magnitude and impact of LQEs on biodiversity and ecosystem resilience

During the LQE, 90 megafauna genera and almost 180 of the largest species of land vertebrates (≥45 kg) disappeared forever from the fossil record (Koch and Barnosky, Reference Koch and Barnosky2006; Barnosky, Reference Barnosky2008; Stuart, Reference Stuart2015). The event is global. It happens over a geologically brief time span of a few tens of thousands of years on all continents, except Antarctica. Nonetheless, there are significant differences in patterns, including the precise timing and severity (Koch and Barnosky, Reference Koch and Barnosky2006; Stuart, Reference Stuart2015). The main commonality is the specific body mass range of the extinct species, with a strong prevalence of large and very large animals. Many weighed hundreds of kilos. Some (for instance, several species of elephantids, South American giant ground sloths and glyptodonts, or the Australian Diprotodon, the largest marsupial ever known) were colossal, exceeding one or more tons in weight. Using Europe as an example, where extinctions were more severe than in Africa but less than in the Americas or Australasia (Stuart, Reference Stuart2015), all terrestrial vertebrates heavier than 1 ton and, overall, more than ¼ of megafauna species, including, among the others, rhinoceros, elephants and mammoths, a giant deer, at least one species of horse, cave lions and cave bears, as well as our own sister lineage, the Neanderthals, vanish at a rate at least 10 times faster than in the previous 7,00,000 years between 50,000 and 10,000 ya (Stuart, Reference Stuart and MacPhee1999). In some cases, like the common hippopotamus, the lion and the spotted hyena, species go locally extinct, leaving behind an impoverished fauna, but they survive in a restricted area of their original range.

LQE often hit keystone species and ecological engineers, whose loss may have triggered cascades of extinctions (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016; Pires, Reference Pires2024). Present-day large predators can have top-down regulatory effects on ecological networks (Ripple et al., Reference Ripple, Estes, Schmitz, Constant, Kaylor, Lenz, Motley, Self, Taylor and Wolf2016; Wilting et al., Reference Wilting, Roellig and Tilker2025). Their presence or absence, or even just a sharp change in population density, can trigger ecosystem shifts that radically alter species composition and the biomass and productivity of an ecological community (Ripple et al., Reference Ripple, Estes, Beschta, Wilmers, Ritchie, Hebblewhite, Berger, Elmhagen, Letnic, Nelson, Schmitz, Smith, Wallach and Wirsing2014, Reference Ripple, Estes, Schmitz, Constant, Kaylor, Lenz, Motley, Self, Taylor and Wolf2016). A textbook example of this type of ecological interaction today is the Pacific sea otter (Estes, Reference Estes2020). Its population density determines whether coastal regions remain in a highly productive, biodiverse kelp forest state, or, when otters are decimated by hunters, predators or disease, they shift to an impoverished sea urchin barren. As reviewed by Malhi et al. (Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016), the largest mega-herbivores also play keystone ecological roles. Elephants promote long-distance large-seed dispersal (reducing inbreeding and allowing plants to track climate change and colonize new regions (Pires, Reference Pires2024) and engineer the environment by creating open habitats, as they reduce woody species cover. A single African elephant can uproot hundreds of adult trees each year, but these animals may also promote the growth of woody vegetation by reducing competition with grasses and by moving nutrients inland from river floodplains, as well as by removing dry biomass and, thus, decreasing the frequency of fires in the savanna. Likewise, in wetter and colder climates, mammoths, horses, woolly rhinos and bisons facilitated the persistence of a steppe habitat at high latitudes. In this case, grasses might have increased transpiration in summer, avoiding upper-soil waterlogging and, therefore, reducing the occurrence of a, nowadays dominant, low-diversity mossy tundra (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016; Pires, Reference Pires2024).

Although the population density of the biggest mega-herbivores is likely primarily controlled by bottom-up factors, such as primary productivity, the largest carnivore species may also contribute through predation on their young (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016). Conversely, if the largest predators significantly preyed on mega-herbivores, the population size of the herbivores could have exerted a bottom-up regulatory influence on the predators (Pires, Reference Pires2024). Furthermore, larger predators directly (by killing) or indirectly (via competition) contribute to regulating the populations and food preferences of smaller megafauna carnivores and meso-predators. As they went extinct, this regulatory effect was lost, and the surviving predators expanded their ecological niche (Pires, Reference Pires2024). Likewise, with fewer predators and without competition from other mega-herbivores, such as, for instance, mammoths and horses, the remaining species of grazers and browsers could increase population density and enlarge their distributional range, as it happened with the bison in North America (Pires, Reference Pires2024). Thus, the megafauna ecosystem is characterized by mutual interactions between several keystone actors, such as the largest primary consumers and predators. These interactions are part of a much larger trophic network, whose complexity is still poorly understood and includes, among others, scavengers and an often overlooked biotic component, that of the megafauna-related microorganisms and parasites (Doughty et al., Reference Doughty, Prys-Jones, Faurby, Abraham, Hepp, Leshyk, Fofanov, Nieto, J-C and Galetti2020; Pires, Reference Pires2024). Clearly, the consequences of LQE went well beyond the loss of many of the most charismatic representatives of the terrestrial fauna (Smith et al., Reference Smith, Smith, Hedberg, Lyons, Pardi and Tomé2023). The local decline or complete disappearance of the majority of the Pleistocene megafauna had disproportionate effects on habitats and ecosystems, which have become impoverished and less functionally redundant, as the functions regulating fluxes of energy and nutrients now depend on fewer species. With loss of functions, or even simply because of a reduced number of animals performing those functions, ecosystems are less resilient and more vulnerable to further future biodiversity losses (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016; Pires, Reference Pires2024).

Indeed, the LQE is the most selective event of the entire Cenozoic (our era, which begins 66 million ya) in terms of body mass of the extinct species, whose average size was between two and three orders of magnitude greater than that of surviving species (Pires, Reference Pires2024). This selectivity alone implies downsizing and ecological downgrading of trophic networks, and, thus, a sharp reduction in landscape heterogeneity and functional diversity (Pires, Reference Pires2024). As an extreme example, the megafauna of South America before the LQE consisted of species of herbivores that were on average larger than their African counterparts (Pires, Reference Pires2024). After the LQE, none of the herbivores heavier than 1 ton survived in South America, and only tapirs and camelids are left among the species with body mass above 100 kg. Thus, as Pires (Reference Pires2024) remarks, today’s Africa has an exceptional richness in very large herbivores, many of which perform important ecosystem functions, but that is mostly an artifact of the devastating effect of LQE in other continents. Earth, after the LQE and before the agricultural and industrial revolution, was already a very different place, one that, for the first time since life recovered from the mass extinction that wiped out all non-avian dinosaurs 66 million ya, was deprived of a diverse, abundant and ecologically functional megafauna (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016).

From megafaunal collapse to planetary crisis: Is human destructiveness a natural trajectory?

LQE happens during the main wave of out-of-Africa dispersal of modern humans (Homo sapiens sapiens). While anthropogenic impacts seem like a likely contributor to the LQE, high-quality data on the subject remain scarce. Archaeological sites associated with megafauna remains are, indeed, rare (Meltzer, Reference Meltzer2015; Nagaoka et al., Reference Nagaoka, Rick and Wolverton2018) and it is often hard to precisely link the arrival of hunter-gatherers with specific extinctions. The Late Quaternary is also a time of rapid changes in climate and habitats, as the last glacial peaks and then ends. Koch and Barnosky (Reference Koch and Barnosky2006) and Stuart (Reference Stuart2015) debate hypotheses and review the evidence, but new studies have been published since then. For instance, two very recent examples, with extensive examinations and sophisticated modeling of the most likely factors, are Stewart et al. (Reference Stewart, Carleton and Groucutt2021) and Lemoine et al. (Reference Lemoine, Buitenwerf and Svenning2023), who reach opposite conclusions, one blaming the climate and the other humans.

Whether human hunters alone (as in the original ‘overkill hypothesis’ (Martin, Reference Martin1966), climate change, other factors or their complex interaction (Koch and Barnosky, Reference Koch and Barnosky2006; Stuart, Reference Stuart2015), have been responsible for the LQE is not the focus of this essay. Having stressed the large uncertainties on causes, I will assume a decisive role of humans in order to discuss the potential implications of the LQE for the modern global environmental crisis. For this assumption to hold, one does not need to postulate a rapid overkill of large animals by humans, as foragers rapidly spread across continents where prey had not evolved effective anti-predator responses to skilled social hunters with relatively sophisticated weapons, including the use of fire. The role of humans is ‘decisive’ even if it was their interaction with rapid changes in the climate, and, thus, with habitat modification, that caused the extinctions. In that case, humans would share the blame with geological forces, but, without us tipping the balance, the Ice Age megafauna would have largely survived environmental change as most of its representatives did in previous glacials and interglacials.

What, then, are the implications of a potentially large-scale extinction event caused by a relatively small number of hunter-gatherers, whose global population at its peak may have been half that of present-day London (Hawks et al., Reference Hawks, Hunley, Lee and Wolpoff2000; Eller et al., Reference Eller, Hawks and Relethford2009), as they emerged from their African homeland toward the end of the Ice Age? The vast majority of LQE occur before there is any radical change in human behavior and ecology (Scerri and Will, Reference Scerri and Will2023). Gradual innovations, better tools and, likely, a progressive development of more complex forms of symbolic communication were facilitated by the demographic expansion of the Upper Paleolithic, but ‘modern human behavior had deep African roots’ (Scerri and Will, Reference Scerri and Will2023, p. 13). Thus, if vast anthropogenic extinctions predate the modern lifestyle of the agricultural and, later, industrial societies, it becomes harder to point at these specific cultural revolutions (O’Brien and Laland, Reference O’Brien and Laland2012) as the root causes of human environmental destructiveness. Nature came before nurture.

If destructiveness is the essence of our species, this begs an important question: ‘Is the human impact natural?’ (Crist, Reference Crist2019, p. 83). In an investigation of the deeper reasons for our current environmental problems (Nelson and Lemberger, Reference Nelson and Lemberger2021), Crist (Reference Crist2019) devotes an entire chapter to this conundrum. The interpretation of the LQE plays a central role in her argument (Crist, Reference Crist2019, pp. 102–104): the LQE ‘… [often] attributed … to the global spread of human hunters … [seems to invite] an irresistible inference … that the anthropogenic extinction crisis is a single continuous event beginning with the human diaspora out of Africa … and continuing all the way to this morning’. Thus, she adds (p. 108–109), ‘the prevalent belief that human nature is to blame for the ecological crisis offers a seductively simple explanation that discourages deeper thinking … naturalizing the human impact on the biosphere … because humans are essentially different from the rest of life’. However, in sharp contrast to this view, her definitive conclusion (p. 109) is that ‘human supremacy is a social achievement … not a biological inheritance’.

The excerpt from Crist’s book provides a useful summary of the seemingly obvious inference of an anthropogenic origin for the LQE, but also of a less apparent, more unsettling, implication: the naturalization of human devastating impacts. Like several other scholars, particularly in the humanities and social sciences (e.g., Deloria [Reference Deloria1997] cited by Sayre [Reference Sayre2001]; Robins [Reference Robins2024]), Crist rejects both the idea of an unlikely continuous pattern of extinctions over the past 50,000 years and, even more strongly, the tendency to naturalize and, therefore, normalize, human dominance over Earth’s ecosystems, which she attributes instead to cultural factors. Similar points have, actually, been made also by natural scientists. For instance, Louys et al. (Reference Louys, Braje, Chang, Cosgrove, Fitzpatrick, Fujita, Hawkins, Ingicco, Kawamura, MacPhee, McDowell, Meijer, Piper, Roberts, Simmons, van den Bergh, van der Geer, Kealy and O’Connor2021, p. 1), in their examination of archaeological and paleontological evidence from Pleistocene islands inhabited by hominins to assess ‘whether humans have always been destructive agents’, concluded that ‘It is not until the Holocene that large-scale changes in technology, dispersal, demography and human behavior visibly affect island ecosystems. The extinction acceleration we are currently experiencing is thus not inherent but rather part of a more recent cultural complex.’

Since I have assumed, for this discussion, that the LQEs were driven by our species, the implication, however, appears inevitable. Human environmental destructiveness is not an existential aspect of the ‘modern lifestyle’ (‘a social achievement’). If this were the case, its emergence should come after, not before, the cultural revolution marked by the progressive domestication of plants and animals, which began during the early millennia of the Holocene and became a global phenomenon between 8,000 and 5,000 ya (Ellis et al., Reference Ellis, Kaplan, Fuller, Vavrus, Klein Goldewijk and Verburg2013; Taiz, Reference Taiz2013; Stephens et al., Reference Stephens, Fuller, Boivin, Rick, Gauthier, Kay, Marwick, Armstrong, Barton, Denham, Douglass, Driver, Janz, Roberts, Rogers, Thakar, Altaweel, Johnson, Sampietro Vattuone, Aldenderfer, Archila, Artioli, Bale, Beach, Borrell, Braje, Buckland, Jiménez Cano, Capriles, Diez Castillo, Çilingiroğlu, Negus Cleary, Conolly, Coutros, Covey, Cremaschi, Crowther, Der, di Lernia, Doershuk, Doolittle, Edwards, Erlandson, Evans, Fairbairn, Faulkner, Feinman, Fernandes, Fitzpatrick, Fyfe, Garcea, Goldstein, Goodman, Dalpoim Guedes, Herrmann, Hiscock, Hommel, Horsburgh, Hritz, Ives, Junno, Kahn, Kaufman, Kearns, Kidder, Lanoë, Lawrence, Lee, Levin, Lindskoug, López-Sáez, Macrae, Marchant, Marston, McClure, McCoy, Miller, Morrison, Motuzaite Matuzeviciute, Müller, Nayak, Noerwidi, Peres, Peterson, Proctor, Randall, Renette, Robbins Schug, Ryzewski, Saini, Scheinsohn, Schmidt, Sebillaud, Seitsonen, Simpson, Sołtysiak, Speakman, Spengler, Steffen, Storozum, Strickland, Thompson, Thurston, Ulm, Ustunkaya, Welker, West, Williams, Wright, Wright, Zahir, Zerboni, Beaudoin, Munevar Garcia, Powell, Thornton, Kaplan, Gaillard, Klein Goldewijk and Ellis2019). Farming dramatically transformed us and the land (Ellis et al., Reference Ellis, Kaplan, Fuller, Vavrus, Klein Goldewijk and Verburg2013; Taiz, Reference Taiz2013). However, if anthropogenic impacts had been devastating the planet well before agriculture, then, a massive ecological footprint becomes a defining trait of human evolution. Thus, not even Paleolithic hunter-gatherers were in harmony with the ecological community they belonged to. As these foragers colonized all nonfrozen continents, they tipped the balance of ecosystems that had, for the most, robustly survived hundreds of thousands of years of dramatic changes in the climate and environment. In this scenario, humans are seen as a consistently hyper-dominant member of the ecological communities for no less than 50,000 years. To put it bluntly, in the light of the consequences of this dramatic dominance, we, as a species, can indeed be seen as ‘destructive by nature’.

Following this line of reasoning, dominant anthropogenic impacts become part of our natural history (‘a biological inheritance’ in Crist’s terms). In the sociological interpretation, an essentialist conclusion becomes an evolutionary fact: ‘This form of species narrative allows people to assume that ecologically destructive behaviors arise from their genetic humanity and not from the cultural and political structures’ (Robins, Reference Robins2024, p. 18). But is it true?

Of course, there are no genes for environmental destructiveness. Natural selection, throughout the history of our lineage, has led to the coevolution of a combination of traits, such as intelligence and sophisticated tool use (including the control of fire), a complex syntactical language that facilitates cooperation and cumulative advances in culture and so on (Richerson et al., Reference Richerson, Gavrilets and de Waal2021). This combination has predisposed our species to an exponential increase in its ability to colonize and successfully survive in almost any type of habitat. Progressively, we also discovered new resources to generate energy and labor, such as domestic species during the Agricultural Revolution and fossil fuels during the Industrial Revolution (Fischer-Kowalski et al., Reference Fischer-Kowalski, Krausmann and Pallua2014). Culture, here defined as the social transmission of information (Bridges and Chittka, Reference Bridges and Chittka2019), was key to progress. Behavioral innovation, tool use and the capacity of rapidly learning from other individuals and groups, and to cooperate in larger numbers (Boyd and Richerson, Reference Boyd and Richerson2024), allowed to flexibly reshape the ecological niche of humans in relation to the specific conditions (Henrich and McElreath, Reference Henrich and McElreath2003). This, in turn, might have led to the preferential selection of gene variants that are favorable in a human-modified environment and, therefore, to a runaway process of accelerated gene-culture coevolution (Laland et al., Reference Laland, Odling-Smee and Myles2010; Rendell et al., Reference Rendell, Fogarty and Laland2011; O’Brien and Laland, Reference O’Brien and Laland2012). As Rendell et al. (Reference Rendell, Fogarty and Laland2011, p. 833) wrote, ‘effective niche-constructing capabilities generate selection for more potent niche-constructing capabilities, in a self-reinforcing dynamic’. For instance, the domestication of animals for meat and dairy farming might have brought a selective advantage to herders and, at the same time, if being able to drink milk after weaning increased fitness, created an environment where lactose tolerance in adults could evolve (Laland et al., Reference Laland, Odling-Smee and Myles2010). Rapid gene-culture coevolution likely played a central role in transforming us into a hyper-keystone species that has a disproportionate impact on ecosystems and affects multiple other keystone species (Worm and Paine, Reference Worm and Paine2016). Usually, our impact is harmful to biodiversity and ecological equilibrium, and it frequently happens on a planetary scale. Thus, as a ‘side effect’ of our own success, we began exhausting resources, leading other life forms to extinction and shifting ecosystems toward degraded states and a precarious balance. Yet, despite local failures, and even occasional societal collapses (Scheffer, Reference Scheffer2016), in the evolutionary short term of a few tens of thousands of years, humanity has managed to escape the consequences of environmental damage, increasing its population from a few million Paleolithic foragers to the present-day 8 billion. However, that the process, as powerful as it is, cannot be sustained indefinitely is clear in Rendell et al.’s words (p. 833): ‘Nonetheless, most resources cannot rise without limit, so at some point an upper bound will be reached, and the opportunities for the evolution of more potent niche construction will diminish’.

If it is a fact that humans are ‘destructive by nature’ (a first implication of the ‘irresistible inference’ of a continuity in anthropic impacts), we have not just discovered an accidental, but rather unfortunate, outcome of our evolutionary history. For the social scientists, naturalizing destructiveness also leads to further and profoundly disturbing corollary claims that can bias, if not undermine, how contemporary society responds to the planetary environmental crisis we face. To begin, since the emergence of modern humans in evolutionary history, true wilderness has never existed. If no human-inhabited environment is, or ever was, pristine, it follows that we all, in every region of the world, regardless of historical socioeconomic and demographic differences, share the responsibility for the anthropogenic devastation of the world’s ecosystems. Besides, even if we are not (and cannot be, from an evolutionary and ecological perspective) separate from nature, a massive ecological footprint defining the human species and its interaction with the environment suggests inevitability: this is part of what we are as a biological species. At best, as humanity confronts the threats of global environmental change, impacts might be mitigated and, if achievable, their directions managed to benefit first and foremost our own species. Thus, any intervention is bound to happen with us, H. sapiens sapiens, as the center stage, as that was and remains the niche our species has evolved to fill.

Change and responsibility: Who bears the burden for the twenty-first-century environmental crisis?

Are we, all living people of all continents and all those who came before us, equally responsible for the anthropogenic environmental crisis of the twenty-first century? The answer to this question is the easiest: we are not, even assuming that Paleolithic hunter-gatherers drove the Ice Age megafauna to extinction, dramatically altering the ecosystem balance on land (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016; Pires, Reference Pires2024). We might be ‘destructive by nature’, in the sense I outlined before. However, there are differences in quality (debatable) and quantity (indisputable).

A few words about quality, first. A human-driven LQE event is a contingent accident. Paleolithic foragers are most unlikely to have intentionally driven any species to extinction. This is in sharp contrast with, for instance, the systematic and meticulous persecution of the thylacine in Tasmania by white farmers, coupled with the destruction of its habitat, until the wild population declined and the last captive individual died (Sleightholme and Campbell, Reference Sleightholme and Campbell2016). Likewise, it is also radically different from the large-scale and deliberate eradication of wolves by European settlers and their descendants in most of North America (Musiani and Paquet, Reference Musiani and Paquet2004).

Besides, we cannot judge indigenous populations that might have eroded biodiversity and degraded ecosystems in the past using contemporary understanding and values (Taylor et al., Reference Taylor, Wieren and Zaleha2016). There is a difference in knowledge, intentionality and awareness. In Best’s words (Best, Reference Best2021, p. 9), ‘extinctions did not always arise from the tips of human spears and orgies of killing but rather one species, among others, exploiting an ecological niche for survival, perhaps unaware of species limits and finitude and the consequences of its actions’. Today, in contrast, a large proportion of humanity is well aware of being the driver of a massive demographic decline in wildlife populations and a myriad of very recent anthropogenic extinctions (Ceballos et al., Reference Ceballos, Ehrlich and Dirzo2017; Cowie et al., Reference Cowie, Bouchet and Fontaine2022). Likewise, pollution and deforestation, just two of many possible examples of environmental degradation, make daily headlines in the media, and we are increasingly learning about the loss of ecosystem functions and the dramatic shift in ecosystem states accompanying the disappearance of keystone species (Malhi et al., Reference Malhi, Doughty, Galetti, Smith, Svenning and Terborgh2016). Yet, aware as we are, we seem unable to stop.

One might argue that, tens of thousands of years ago, as today, ours is the only species able to drive other life forms to extinction on a geological scale (Ceballos et al., Reference Ceballos, Ehrlich and Dirzo2017; Cowie et al., Reference Cowie, Bouchet and Fontaine2022). Wherever we have been, we seem to have left evidence of our devastating ecological footprint (Denevan, Reference Denevan1992; Foley et al., Reference Foley, Gronenborn, Andreae, Kadereit, Esper, Scholz, Pöschl, Jacob, Schöne, Schreg, Vött, Jordan, Lelieveld, Weller, Alt, Gaudzinski-Windheuser, Bruhn, Tost, Sirocko and Crutzen2013; Ellis et al., Reference Ellis, Gauthier, Klein Goldewijk, Bliege Bird, Boivin, Díaz, Fuller, Gill, Kaplan, Kingston, Locke, McMichael, Ranco, Rick, Shaw, Stephens, J-C and Watson2021). There might be a few exceptions, if any. According to a Cherokee scholar, even ‘modern Natives and their ancestors are neither saints nor sinners in environmental matters – sometimes they have degraded their environments even to the point of societal collapse’ (Weaver [Reference Weaver1996, p. 7] cited by Taylor et al. [Reference Taylor, Wieren and Zaleha2016]). However, whether or not one agrees that the core of who we are and how we interact with other species and the environment has changed little over the past 50,000 years, the quantitative differences in impacts between then and now are undeniable. These are measurable. A few examples should suffice to remove doubts on magnitude and responsibility. Let us focus primarily on assessing biodiversity and environmental impacts before and after the Industrial Revolution, a period for which we have uncontested evidence. If it can be demonstrated that, despite a long history of widespread extinctions and global impacts, the decline of wild species and habitat degradation was relatively modest before the industrial era, the assignment of responsibility becomes clear: the blame lies with the dominant industrialized nations.

The majority of scholars acknowledge the exponential escalation in anthropogenic impacts in the last few centuries, and a further acceleration after World War II (Crutzen, Reference Crutzen, Ehlers and Krafft2006; Foley et al., Reference Foley, Gronenborn, Andreae, Kadereit, Esper, Scholz, Pöschl, Jacob, Schöne, Schreg, Vött, Jordan, Lelieveld, Weller, Alt, Gaudzinski-Windheuser, Bruhn, Tost, Sirocko and Crutzen2013; Smith and Zeder, Reference Smith and Zeder2013; Steffen et al., Reference Steffen, Broadgate, Deutsch, Gaffney and Ludwig2015; Lewis and Maslin, Reference Lewis and Maslin2020). Crist (Reference Crist2019) herself writes (p. 107) that ‘humanity’s blow on the biosphere has increased by orders of magnitude in comparison to what occurred previously in the Holocene’. Accordingly, Crutzen (Reference Crutzen, Ehlers and Krafft2006, p. 14) offers a number of examples to support his proposal of a new geological epoch, the Anthropocene, marked by a massive planetary increase in anthropic influence: ‘During the past … centuries human population increased tenfold to 6000 million, growing by a factor of four during the past century alone … accompanied for example, by a growth in the cattle population to 1400 million … Similarly … industrial output … grew forty times. More than half of all accessible fresh water is used by humankind. Fisheries remove more than 25% of the primary production of the oceans in the upwelling regions and 35% in the temperate continental shelf regions’. Now, two decades after Crutzen’s proposal, an even larger population and an additional increase in global consumerism have further magnified those impacts (Fletcher et al., Reference Fletcher, Ripple, Newsome, Barnard, Beamer, Behl, Bowen, Cooney, Crist, Field, Hiser, Karl, King, Mann, McGregor, Mora, Oreskes and Wilson2024). For instance, CO2 is more than 50% higher than in preindustrial time and still on the rise (Friedlingstein et al., Reference Friedlingstein, O’Sullivan, Jones, Andrew, Bakker, Hauck, Landschützer, Le Quéré, Luijkx, Peters, Peters, Pongratz, Schwingshackl, Sitch, Canadell, Ciais, Jackson, Alin, Anthoni, Barbero, Bates, Becker, Bellouin, Decharme, Bopp, Brasika, Cadule, Chamberlain, Chandra, Chau, Chevallier, Chini, Cronin, Dou, Enyo, Evans, Falk, Feely, Feng, Ford, Gasser, Ghattas, Gkritzalis, Grassi, Gregor, Gruber, Gürses, Harris, Hefner, Heinke, Houghton, Hurtt, Iida, Ilyina, Jacobson, Jain, Jarníková, Jersild, Jiang, Jin, Joos, Kato, Keeling, Kennedy, Klein Goldewijk, Knauer, Korsbakken, Körtzinger, Lan, Lefèvre, Li, Liu, Liu, Ma, Marland, Mayot, McGuire, McKinley, Meyer, Morgan, Munro, Nakaoka, Niwa, O’Brien, Olsen, Omar, Ono, Paulsen, Pierrot, Pocock, Poulter, Powis, Rehder, Resplandy, Robertson, Rödenbeck, Rosan, Schwinger, Séférian, Smallman, Smith, Sospedra-Alfonso, Sun, Sutton, Sweeney, Takao, Tans, Tian, Tilbrook, Tsujino, Tubiello, van der Werf, van Ooijen, Wanninkhof, Watanabe, Wimart-Rousseau, Yang, Yang, Yuan, Yue, Zaehle, Zeng and Zheng2023), but climate change is just one of many critical aspects of the present global environmental crisis: six (biosphere integrity, changes in land and freshwater systems, alteration of biogeochemical cycles of nutrients, etc.) of nine planetary boundaries for a safe operating space for humanity have been crossed (Richardson et al., Reference Richardson, Steffen, Lucht, Bendtsen, Cornell, Donges, Drüke, Fetzer, Bala, von Bloh, Feulner, Fiedler, Gerten, Gleeson, Hofmann, Huiskamp, Kummu, Mohan, Nogués-Bravo, Petri, Porkka, Rahmstorf, Schaphoff, Thonicke, Tobian, Virkki, Wang-Erlandsson, Weber and Rockström2023) and we rapidly approximate tipping points of irreversible change (Lenton et al., Reference Lenton, Rockström, Gaffney, Rahmstorf, Richardson, Steffen and Schellnhuber2019; Tollefson, Reference Tollefson2023).

That the Industrial Revolution has been a multiplier of the ecological footprint of humanity, as never before, can be appreciated also by measuring the massive global reduction in wildlands, the steep increase in the rate of extinction and the different magnitude of the sociometabolism of foragers, farmers and industrialized societies. Using a definition of wildlands as uninhabited regions under no intense use, Ellis et al. (Reference Ellis, Klein Goldewijk, Siebert, Lightman and Ramankutty2010) conclude (p. 589) that ‘in 1700, nearly half of the terrestrial biosphere was wild … [and] most of the remainder was in a seminatural state (45%) having only minor use for agriculture and settlements. By 2000, the opposite was true, with the majority of the biosphere in agricultural and settled anthromes [i.e., anthropogenic biomes], less than 20% seminatural and only a quarter left wild’. Wilderness has, thus, been vanishing before our eyes in the last 250 years. ‘Wild’ does not mean ‘untouched’. Prehistoric activities by humans might have reshaped habitats and ecological communities, leaving a long-term legacy on the biosphere (Ellis et al., Reference Ellis, Gauthier, Klein Goldewijk, Bliege Bird, Boivin, Díaz, Fuller, Gill, Kaplan, Kingston, Locke, McMichael, Ranco, Rick, Shaw, Stephens, J-C and Watson2021). However, the strongest impacts, responsible for the current crisis, are a consequence of the direct or indirect appropriation of land and the intensification in the use of resources by industrialized societies (Ellis et al., Reference Ellis, Klein Goldewijk, Siebert, Lightman and Ramankutty2010, Reference Ellis, Kaplan, Fuller, Vavrus, Klein Goldewijk and Verburg2013, Reference Ellis, Gauthier, Klein Goldewijk, Bliege Bird, Boivin, Díaz, Fuller, Gill, Kaplan, Kingston, Locke, McMichael, Ranco, Rick, Shaw, Stephens, J-C and Watson2021). Likewise, for millennia we have been responsible for species extinctions on islands and continents (Johnson et al., Reference Johnson, Balmford, Brook, Buettel, Galetti, Guangchun and Wilmshurst2017). Yet, it is in the last three centuries that the rate of vertebrate extinctions has risen to no less than 100 times higher than the background rate estimated using the fossil record (Ceballos et al., Reference Ceballos, Ehrlich, Barnosky, García, Pringle and Palmer2015). This looks impressive, but population declines and species loss are underestimated, as we lack accurate information on the past, and even today, only a minority of taxa is carefully assessed by conservationists (Hughes et al., Reference Hughes, Qiao and Orr2021). Thus, it might not be unreasonable that, since 1,500 AD, 8–13% of the currently known 2 million species have been lost (Cowie et al., Reference Cowie, Bouchet and Fontaine2022) and, as in land vertebrates (Ceballos et al., Reference Ceballos, Ehrlich, Barnosky, García, Pringle and Palmer2015), this has likely happened with an accelerated pace in the last few 100 years.

In the attribution of the responsibility of anthropic impacts, however, the type of economy and the prevalent societal lifestyle (a term here used, for brevity, to broadly refer to how people live their lives (Agnew et al., Reference Agnew, Pettifor and Wilson2023) matter. In calculating the human ecological footprint (Wackernagel et al., Reference Wackernagel, Onisto, Bello, Callejas Linares, Susana López Falfán, Méndez Garcı́a, Isabel Suárez Guerrero and Guadalupe Suárez Guerrero1999; Toth and Szigeti, Reference Toth and Szigeti2016), consumption is a key factor and also, often, the largest contributor to species extinction risk (Irwin et al., Reference Irwin, Geschke, Brooks, Siikamaki, Mair and Strassburg2022). For instance, the overconsumption of meat products in the European Union (EU) is linked to the destruction of the Amazon forest and its biodiversity. Estimates (Rajão et al., Reference Rajão, Soares-Filho, Nunes, Börner, Machado, Assis, Oliveira, Pinto, Ribeiro, Rausch, Gibbs and Figueira2020) suggest that ~20% of Brazilian soy, mostly imported in Europe as cattle feed, and almost 50% of beef from Brazil, which provides up to 40% of EU imports, come from illegally deforested areas in this country. The destruction caused to tropical forests around the world by consumption in Europe is such that it has been argued that the ‘Europe’s Green Deal offshores environmental damage to other nations’ (Fuchs et al., Reference Fuchs, Brown and Rounsevell2020, p. 671).

Measuring societal consumption is, thus, important to appreciate direct and indirect anthropic impacts and, therefore, differences in responsibility. A useful concept to convey the magnitude of variation in the consumption of energy and goods during the major transitions from hunter-gatherers to farmers and from the latter to the modern industrialized economy is the metabolic rate of a society or its ‘sociometabolism’. The sociometabolism represents ‘the rate at which energy is exchanged between a human society and its environment, and transformed within a society’ (Malhi, Reference Malhi and Spencer2014, p. 144). With this type of measure, we can estimate the overall material and energy requirement of a person within a given type of society, as if the society were a ‘super-organism’ with a metabolism. Thus, expressed in kcal per day, an average hunter-gatherer needs ~6,000 kcal/day, which is three times the 2,000 kcal/day physiological basal metabolism of a person at rest (Malhi, Reference Malhi and Spencer2014). In contrast, a European preindustrial farmer used about 42,000 kcal/day, and a citizen of a highly developed nation, such as the United States, consumes almost 250,000 kcal/day (Malhi, Reference Malhi and Spencer2014). This means that, in terms of natural resource use, the average American uses as much as 40 hunter-gatherers or six preindustrial European farmers, with each of those farmers using as much energy as seven Paleolithic foragers. Thus, Fischer-Kowalski et al. (Reference Fischer-Kowalski, Krausmann and Pallua2014, p. 8) observe that ‘until … [around AD 1500], human population growth and metabolic rates carry about equal weight in increasing human pressure on the environment approximately fivefold from the year AD 1 onwards. From then on, the overall pressure of humanity upon the Earth increases by one order of magnitude; energy intensity contributes to this rise by roughly tripling the impact of population growth. Technology, because it is based upon a shift from biomass to fossil fuels … does not moderate this impact, but enhances it by a factor of 1.5’. They are also clear (p. 25) that it is with the incipient Industrial Revolution that ‘a veritable take-off can be observed. From 1700 onwards, human impact doubles every century, from 1900 on it doubles in 50 years, and from 1950 on it triples in 50 years, with no sign of saturation yet’. This clearly shows that the biggest rise in human impact on the environment, although already noticeable a couple of centuries before industrialization began, is recent, consistent with previously mentioned indicators such as extinction rates, population growth, CO2 emissions and the amount of wildlands.

With developing countries having a sociometabolism 5–10 times lower than that of industrialized nations, we arrive at a less obvious conclusion: in modern times, the impact of the socioeconomic system far outweighs that of population growth (Krausmann et al., Reference Krausmann, Fischer-Kowalski, Schandl and Eisenmenger2008). The direct implication is that societies that first transitioned to the industrial system have historically contributed the most to environmental damage – about 10 times more than farming societies and roughly 50 times more than hunter-gatherer societies (Krausmann et al., Reference Krausmann, Fischer-Kowalski, Schandl and Eisenmenger2008; Fischer-Kowalski et al., Reference Fischer-Kowalski, Krausmann and Pallua2014). These industrialized nations continue to have a disproportionate ecological footprint, despite smaller populations, when compared to developing countries and less wealthy societies. Thus, going back to greenhouse gas emissions as an example of impact, the wealthiest 10% of the world population (mainly the ~700 million citizens of Europe and North America) was responsible in 2015 for 48% of the total annual emissions, while the 50% bottom income earners (the poorest ~3.7 billion people) emitted overall just 7% of those same atmospheric pollutants (Capstick et al., Reference Capstick, Khosla and Wang2020). However, it is not just current emissions that are driven by rich, industrialized countries. Since 1800, 86% of the contribution to the total increase of CO2 in the atmosphere has been attributed to highly developed countries (Neumayer, Reference Neumayer2000). It seems, at a first glance, paradoxical that precisely by using the energy of fossil fuels we have managed to increase today’s total mammal biomass of the land megafauna about seven times compared to its estimate before the time of the LQE (Barnosky, Reference Barnosky2008). However, even including marine mammals, and thus the whales and the other giants among living animals, 96% of the total modern-day mammalian biomass is made of humans and their livestock, and just 4% is wild species (Bar-On et al., Reference Bar-On, Phillips and Milo2018). This massively unbalanced ratio indicates a similarly unbalanced ecosystem and is another feature of modernity, since for most of the Pleistocene, and even at the time of the major collapse of the Ice Age megafauna, the human biomass was a fraction of the total biomass of all land mammals (Barnosky, Reference Barnosky2008).

Again and again, we have demonstrated that the ecological footprint of modernity is incomparably heavier than any damage caused in our hunter-gatherer past. Besides, since the spread of agriculture, impacts have been magnified multiple times, and later, with the Industrial Revolution, they have further increased by at least an order of magnitude. For this last and most substantial rise in anthropogenic pressure, the responsibility can be attributed for the vast majority to the highly industrialized Western nations that have so far received the greatest benefit (e.g., the proportion of contribution to the CO2 concentration explains 2/3 of 1990 differences in per capita gross national product (Neumayer, Reference Neumayer2000) to the detriment of the poorest and the environment. In terms of extinctions, the differential contribution is so remarkable that, by quantifying the consequences of consumption on biodiversity, researchers can show how virtually all rich industrialized European and North American countries are, in fact, importing the extinction risk (i.e., causing extinctions elsewhere) from the developing nations, whose resources they are exploiting (Irwin et al., Reference Irwin, Geschke, Brooks, Siikamaki, Mair and Strassburg2022).

To summarize, human-driven LQE cannot be used to argue a shared responsibility of humanity through time for the global environmental damage and the unbalanced ecosystem we have created. In fact, even if we narrowly focus on modern impacts caused by industrialized societies, by far the largest of any time, both currently and historically, the responsibility mostly rests on the shoulders of the richest people and nations. Anthropogenic LQE, therefore, do not make all humans and socioeconomic systems equally impactful: as we may have to acknowledge an early origin of human ‘destructiveness’, we must also recognize that an ecological footprint so huge to alter the climate and cause a rate of extinctions comparable to that of the past five great mass extinctions of geological origin is an unprecedented emerging aspect of the last 250 years of Western-dominated industrialized economy. If anything, clarifying similarities and differences between human-driven extinction events and the related large-scale loss of ecosystemic balance contributes to understanding how the massive inequality within and among nations in the more recent history of humanity is at the base of the current environmental emergency and, thus, a priority to address for any effective solution to the planetary crisis (Ross, Reference Ross2017; Best, Reference Best2021; Fletcher et al., Reference Fletcher, Ripple, Newsome, Barnard, Beamer, Behl, Bowen, Cooney, Crist, Field, Hiser, Karl, King, Mann, McGregor, Mora, Oreskes and Wilson2024).

Rethinking the myth of human incompatibility with balanced ecosystems

Even without any ‘genes for destructiveness’, and even when differences in the magnitude and responsibility of our past and present impacts are acknowledged, one might still wonder whether an anthropogenic LQE ultimately implies a fixed trajectory of human behavior, one that leaves no alternative but to hopelessly document the ongoing destruction of life on Earth. Alternatively, as Nagaoka et al. (Reference Nagaoka, Rick and Wolverton2018) argued, ‘… like any other organism, humans can destroy, modify, enhance, or preserve depending on context. And there is an extensive continuum of human–environment interactions that range from extinctions to sustainable coexistence’ (Nagaoka et al., Reference Nagaoka, Rick and Wolverton2018, p. 9692). Thus, for these scholars, in their evaluation of how different disciplines interpret the overkill model, a major conclusion is that there is no inevitability in human destructiveness as long as we can profoundly reshape our niche in a direction different from the one prevailing since the Agricultural Revolution. Who is right?

In this last section, I will further explain why Nagaoka et al. are correct and argue that, regardless of the onset of widespread anthropogenic impacts, human nature offers no excuse for inaction in the face of an increasingly threatening global environmental crisis. Indeed, we may be ‘destructive by nature’ with a long chain of anthropogenic extinctions that begins 50,000 ya, if not earlier. However, as anticipated, the most profound changes for us and the planet are much more recent and mostly driven by culture. First, the domestication of animals and plants and, later, the industrialization of production and society demonstrate the enormous potential consequences of a culturally dominated niche construction in humans. Regrettably, these revolutions, as they led us to modern civilization and ‘progress’, have also created the unprecedented environmental crisis that threatens our own future survival (Wright, Reference Wright2004). They transformed H. sapiens sapiens in a geological force of destruction that can change the face of the planet to meet its exclusive short-term needs (Ehrlich and Ehrlich, Reference Ehrlich and Ehrlich2008). The root of the current crisis may well be that ‘the will to dominate exists within us in seed form, affiliated perhaps with life’s imperative for self-preservation’ (Crist, Reference Crist2019, p. 112) but it is ‘the anthropocentric worldview’ that has tilled and fertilized ‘that particular seed, growing it into a … now global superstructure of domination’. Much like a positive feedback loop, cultural innovations led to progress, which increased human dominance. This, in turn, tends to reinforce anthropocentrism, promoting the idea of human supremacy and further domination. Over time, this dynamic spiraled out of control, reaching its apex with the ubiquitous Western socioeconomic system of the capitalist free market. To be fair, although neoliberal capitalism currently dominates with devastating impacts, communism and socialism had no better environmental record (Change, Reference Change2011).

Having acknowledged the overwhelming responsibility of industrialized nations in the modern crisis, one might also note that if no totally pristine habitat remains on Earth, and if anthropogenic impacts extend back to historical or even prehistorical times, then, despite the diversity of human behaviors and the differences in socioeconomic structures and belief systems, there appear to be few examples of human societies whose footprint on the environment and biodiversity was truly negligible (Foley et al., Reference Foley, Gronenborn, Andreae, Kadereit, Esper, Scholz, Pöschl, Jacob, Schöne, Schreg, Vött, Jordan, Lelieveld, Weller, Alt, Gaudzinski-Windheuser, Bruhn, Tost, Sirocko and Crutzen2013; Boivin et al., Reference Boivin, Zeder, Fuller, Crowther, Larson, Erlandson, Denham and Petraglia2016; Hussain and Baumann, Reference Hussain and Baumann2024). Nonetheless, humans have achieved an undeniably impressive demographic success, and on average, we also live longer and more comfortable lives. This reading of human history is, however, misleadingly myopic (Wright, Reference Wright2004). Human success is measured on a minimalistic evolutionary timescale of centuries or millennia, despite looming preoccupation and uncertainty about our own future in the next decades (Gross, Reference Gross2013; Best, Reference Best2021; Gross, Reference Gross2023b). Furthermore, the benefits of ‘progress’ were and are highly unequally distributed, and originate from colonialism and economic imperialism (Ross, Reference Ross2017; Lewis and Maslin, Reference Lewis and Maslin2020; Ghosh, Reference Ghosh2021; Hickel et al., Reference Hickel, Dorninger, Wieland and Suwandi2022) that can be seen as an intraspecific variant of supremacism. A just system should have already eradicated poverty and provided all of humanity with its basic needs. Furthermore, innovations have nearly constantly led to a progress trap, where damage is either externalized (i.e., inflicted on those that did not cause it) or requires further innovation to be fixed (Wright, Reference Wright2004; Ellis et al., Reference Ellis, Kaplan, Fuller, Vavrus, Klein Goldewijk and Verburg2013; Lewis and Maslin, Reference Lewis and Maslin2020; Best, Reference Best2021). This is how, for instance, the slow progressing and often problematic (Bell, Reference Bell2015; Sovacool, Reference Sovacool2021; Gross, Reference Gross2024) ‘ecological transition’ of wealthier industrialized nations outsources social and environmental damage to developing countries by exploiting their natural resources (Fuchs et al., Reference Fuchs, Brown and Rounsevell2020), as with Chilean copper or Congolese coltan and cobalt used in low emission electric vehicles (Sovacool et al., Reference Sovacool, Ali, Bazilian, Radley, Nemery, Okatz and Mulvaney2020). Likewise, the supraspecific application of the same supremacist philosophy is wiping out thousands of species, destroying habitats, polluting air, soil and water and degrading and downgrading ecosystems (Crist, Reference Crist2019; Ghosh, Reference Ghosh2021). Environmental impact interacts with social injustice, as it hits harder the poorest among people and nations, ironically, precisely those who contributed the least to the crisis (Chancel, Reference Chancel2022; Fletcher et al., Reference Fletcher, Ripple, Newsome, Barnard, Beamer, Behl, Bowen, Cooney, Crist, Field, Hiser, Karl, King, Mann, McGregor, Mora, Oreskes and Wilson2024; Wilkinson and Pickett, Reference Wilkinson and Pickett2024).

Transitioning to a radically new socioeconomic system is challenging and faces resistance (Büchs and Koch, Reference Büchs and Koch2019), but these difficulties do not originate from a potentially inherent destructiveness of humans. They derive from the structural quality and complexity of an ill-designed ecological niche constructed to achieve human dominance with no concern for ecosystem balance and other life forms (Büchs and Koch, Reference Büchs and Koch2019). Yet, the direction for change is clear: we are part of the Earth ecosystem, and a stable economy and thriving society depend on its long-term health and robustness (Ruuska et al., Reference Ruuska, Heikkurinen and Wilén2020; Dirzo et al., Reference Dirzo, Ceballos and Ehrlich2022; Fletcher et al., Reference Fletcher, Ripple, Newsome, Barnard, Beamer, Behl, Bowen, Cooney, Crist, Field, Hiser, Karl, King, Mann, McGregor, Mora, Oreskes and Wilson2024). The United Nations Sustainable Development Goals (SDGs) have, so far, been largely contradictory and ineffective (Hickel, Reference Hickel2019; Cardini, Reference Cardini2024; Montesano Montessori and Lautensach, Reference Montesano Montessori and Lautensach2024). However, they do present, at least in theory, a realistic vision of priorities (Montesano Montessori and Lautensach, Reference Montesano Montessori and Lautensach2024), as illustrated by the SDG ‘wedding cake’ model (https://youtu.be/dcvz6Fv8DqU), where the biosphere forms the broad foundational layer supporting society, which in turn underpins the economy.

The ecological niche we have constructed rests on a no longer tolerable supremacist assumption of inequality. This niche is part of our phenotype, a complex manifestation of the interaction between our genes and the environment. However, it is a niche that, in H. sapiens sapiens, has been largely dominated by culture. Genetics may set the direction of our behavior, but the final outcome, what we end up doing, depends on the specific environment, which in humans is strongly shaped by culture. Thus, even the most fundamental instinct, without which there can be no evolution by natural selection, that is, the desire to survive and reproduce, is not destiny (Alcock, Reference Alcock2001). Many people choose to have few or no children at all. From an evolutionary standpoint, this is a contradiction unless it somehow benefits one’s relatives, like in the concept of kin selection (Birch, Reference Birch2019). While this can happen in some cases, it is probably uncommon in modern societies. Surely, we did not choose either our genes or the environment (Sapolsky, Reference Sapolsky2021) that led us to this ‘decision’, but the mere fact that, given a specific context, the outcome goes against the strongest genetic predisposition, that of leaving a progeny, shows that genes, our ‘essence’, are not fate (p. 182 and following ones in Alcock [Reference Alcock2001]).

Similarly, while our species may have become a hyper-keystone species (Worm and Paine, Reference Worm and Paine2016) capable of dominating all other components of its ecosystem, this does not mean we must do so. On the contrary, acknowledging this potential can help us transform dominance into a balanced, ecosystemic coexistence, an essential step for ensuring our long-term evolutionary survival as part of a diverse and resilient ecological community (Washington et al., Reference Washington, Taylor, Kopnina, Cryer and Piccolo2017). We are masters of niche construction and ‘niche construction is frequently influenced by prior selection, but it is, at most, only partly determined by genes’ and largely depends ‘on environmental influences and on other inherited information and materials that together shape the developing organism and fashion how it interacts with the world’ (O’Brien and Laland, Reference O’Brien and Laland2012, pp. 436–437). Even if our natural history had made us uniquely able to change the planet with little oversight for the future, it did not remove us from the long-term reliance on the global ecosystem, whose functions go well beyond providing short-term services for humans (Washington et al., Reference Washington, Taylor, Kopnina, Cryer and Piccolo2017). Indeed, our epoch ‘ought to be characterized not by even further distending human hubris, but by humility and respect’, as we ‘cannot ignore the weight of the arguments that provide compelling reason to believe that at least some parts of nonhuman [life] have intrinsic value, and therefore deserve direct moral consideration’ (Batavia and Nelson, Reference Batavia and Nelson2017, p. 374).

Finally, when examining the potential implications of the specific pattern of human-driven extinctions through time for both the attribution of responsibility and the direction we choose to take in our relationships with the biotic and abiotic components of the world ecosystem, we should start from the same fundamental premise of sociobiological research (Segerstråle, Reference Segerstråle2000, p. 389): ‘The problem is … the idea that our moral judgments ought somehow to be tied to the latest scientific knowledge [but]… it is exactly this connection between scientific understanding and perceived social utility that needs to be broken, not least for the reason that scientific knowledge and the interpretation of facts changes over time’. Thus, whether we truly are ‘destructive by nature’ and what precisely this means are elements of a complex and dynamic system of scientific investigation that constantly revises its results. This system provides useful information in the realm of ethics and behavior, but it offers neither a prescription nor a justification for how we should act. Human destructiveness may be the outcome of an evolutionary history that made us the indisputable champion of niche construction. Dominance and supremacism are problematic parts of the niche we built. As we created this ecologically distorted worldview, we can also learn to deconstruct these devastating aspects of our culture to rebalance the human niche within the planetary ecosystem. There is no essentialist obstacle to such a sharp change, and there might be a multiplicity of ways to shift from an anthropocentric to an ecosystem-centered direction. Whether in practice this is feasible rapidly enough to counter the impending environmental disaster is, however, a much harder question to answer.

The first modern humans who colonized the Americas might have caused extinctions and environmental change. That there, and in other continents, a tiny population of ‘low-tech’ foragers was potentially able to irreversibly and radically disrupt the balance of entire ecosystems, invites profound considerations on the sustainability of the human enterprise in the twenty-first century. To what extent LQE, if human-driven, was an accident or might have occurred with a degree of intentionality is hard to say. Most likely, Paleolithic hunter-gatherers were initially unaware of the consequences of their impact. Unlike them, in sharp contrast, but similarly to General Carleton, we know we are leading other life forms to extinction and can clearly see the environmental devastation we bring and the perils ahead of us. Tragically, we may lack a deep appreciation of the urgency and gravity of the crisis. Yet, the end of human dominance should be, for us, the beginning of the next ecocentric cultural revolution of H. sapiens sapiens.

Open peer review

To view the open peer review materials for this article, please visit http://doi.org/10.1017/ext.2025.10007.

Acknowledgements

The author is grateful to Eileen Crist for the brief discussion they had on the implications of the LQE, which contributed to the author’s motivation to further explore an issue that the author has been thinking about for years and became even more interested in after reading her thought-provoking book Abundant Earth. The author would also like to thank the editors, as well as two anonymous reviewers, for their comments, which helped the author improve the article by better stressing its focus and implications. This article is dedicated to the memory of Maria Zaytsava (2001–2025) for her idealism and courage against injustice and the domination of the powerful.

Author contribution

The author designed and wrote the whole study.

AI-assisted technologies (ChatGPT 4) were used only to improve English. The author has reviewed and takes responsibility for the final text.

Financial support

There was no financial support.

Competing interests

The author declares none.

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Author comment: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R0/PR1

Comments

Dear Editor,

please, find submitted a review paper entitled:

Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis.

In the manuscript, I am exploring the broader implications of a possible human role in the Late Quaternary extinction, which led to the loss of more than half of the land megafauna. The paper integrates scientific and humanistic perspectives to argue that human-driven global change may be a fundamental aspect of the interaction between biological and cultural evolution in our species. However, I contend that this trajectory is not predetermined by evolution: we have the capacity to shift our impact from harmful to beneficial.

The review is aimed at a broad readership at the boundary between science and humanities.

If allowed by the journal policy, I suggest a few preferred reviewers:

Eileen Crist, Virginia Tech, Blacksburg VA: ecrist@vt.edu

Anthony Barnosky, UC Berkeley: barnosky@berkeley.edu

Jens-Christian Svenning, Aarhus Universitet: svenning@bio.au.dk

I look forward to your response, and thank you for considering my work.

Sincerely

Andrea Cardini

Review: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R0/PR2

Conflict of interest statement

Reviewer declares none.

Comments

From the abstract, the manuscript appears to aim to challenge the notion that humans are inherently destructive and instead emphasize the capacity of humans to bring about positive change. However, the manuscript’s structure and argumentation do not clearly follow this path. Instead, the first half of the manuscript focuses on Late Quaternary extinctions (LQE) as evidence of human destructiveness, eventually taking the stance that this trait is a given. The manuscript then proceeds to describe modern environmental impacts associated with human actions, identifying mass consumption and exponential population growth as significant factors contributing to the current ecological situation. The conclusion appears to be that human “culture” offers a means to address the environmental degradation that is occurring today. However, the consequence of this argument structure is that the link between the supposed human-caused LQE extinctions and modern environmental degradation breaks down. This leaves the manuscript feeling like it is telling two separate stories.

The conceptual fingerprints of the overkill model are found throughout the manuscript. Unfortunately, the use of overkill in the argument automatically forces the discussion of human impacts to destructiveness as an inherent and fixed trait. However, the second half of the manuscript argues for the ability of humans, through culture, to overcome environmental challenges. In other words, the first half is based on the argument of the fixed nature of human behavior, while the second half leans into the idea that human behavior can strive for solutions. To be logically consistent, the manuscript could retain the LQE premise with its fixed human behavior and dedicate the second half to documenting modern destruction, with no solutions or hope. Alternatively, the manuscript could be reoriented to shift the prehistoric narrative to focus on human behavior as situationally diverse and creative to support the potential for humans to mitigate environmental conditions.

Much of what is described in the second section on the impacts of LQE would have occurred whether it was caused by climate change or human predation.

The quotes and examples referring to indigenous peoples should be removed (page 3, line 43; page 8, line 40; page 11, line 14). The first paragraph suggests that the concept of “survival of the fittest” justifies destructive colonial actions. Given the positionality of Western research relative to indigenous peoples in colonial contexts, we should be thoughtful about using their words and experiences in our argumentation.

Much of the critique of modern societies, such as “the will to dominate” (page 15, line 52) and supremacy over nature, is related to Western ideologies rather than a global trait of humans. This discussion itself could further support the argument that human beliefs and actions vary. However, as it stands, it could be used to support the fixed nature of humans.

There is an argument that group-level selection may occur within humans when we think of solutions related to mitigating the tragedy of the commons. Laws, policies, and cultural norms become a means of constraining individual behavior to benefit the group. Thus, it could support a way for humans to reduce their environmental impact and become more a part of nature. One thing that humans do that stands out is that we excel at creating a great deal of intra-generational phenotypic variability for evolutionary processes to play around with. To constrain our views on human potential as fixed and hard-wired seems like a limiting belief we should avoid.

Review: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R0/PR3

Conflict of interest statement

Reviewer declares none.

Comments

This sort of philosophical musing about the long-term meaning or implications of events and processes in the past does not resolve issues raised in the text. There are no empirical facts, no testing of hypotheses, no falsifying of hypotheses – and therefore no scientific support for either of the polarized positions implied in the manuscript – “anthropomorphic supremacy” (evolution of inevitable human destructiveness) or something else (to be aspired to, such as changing human views towards restoring/conserving ecosystems). Authoritarian religions and hyper-capitalism direct humans to exploit their supremacy, which prehistoric Homo sapiens also might have been in the process of accepting, especially when globally expanding their range.

Recommendation: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R0/PR4

Comments

Both reviewers raised concerns with the manuscript. Reviewer 1 finds the two parts of the paper contradictory and suggests they be better integrated, while Reviewer 2 notes that you provide limited evidence to support either position. Please pay careful attention to Reviewer 1’s suggestions for reframing the manuscript to resolve this contradiction. To address Reviewer 2’s commentary, the argument would be strengthened by incorporating additional empirical support.

Decision: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R0/PR5

Comments

No accompanying comment.

Author comment: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R1/PR6

Comments

Milan October 2025

Dear Editors,

I am grateful to you and the reviewers for considering my contribution and for the constructive comments.

I am answering all questions below (in green), and have tracked changes to the manuscript, as suggested. I also uploaded a clean version with all changes accepted.

Changes were small, as suggested in your letter, but they have definitely helped to clarify potential misinterpretations and better stress why human ‘nature’ provides no excuse for passively accepting anthropogenic impacts and, thus, resigning to the inevitability of the global environmental crisis.

I look forward to hearing from you in due time.

Sincerely

Andrea Cardini

Dipartimento di Scienze Chimiche e Geologiche, Università di Modena e Reggio Emilia, Via Campi, 103 - 41125 Modena – Italy

School of Anatomy, Physiology and Human Biology, The University of Western Australia, 35 Stirling Highway, Crawley WA 6009, Australia

E-mail address: alcardini@gmail.com, andrea.cardini@unimore.it, andrea.cardini@uwa.edu.au

Tel. 0039 059 2058472

https://orcid.org/0000-0003-2910-632X

---------- Forwarded message ---------

From: Cambridge Prisms: Extinction <onbehalfof@manuscriptcentral.com>

Date: Tue, 14 Oct 2025 at 00:08

Subject: Cambridge Prisms: Extinction - Decision on EXT-2025-0018

To: <andrea.cardini@uwa.edu.au>, <andrea.cardini@unimore.it>

Cc: <andrea.cardini@uwa.edu.au>, <andrea.cardini@unimore.it>

13-Oct-2025

Dear Dr. cardini,

EXT-2025-0018 entitled "Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis." which you submitted to Cambridge Prisms: Extinction, has been reviewed. The comments of the reviewer(s) are included at the bottom of this letter. The reviewer(s) have recommended publication, but also suggest some minor revisions to your manuscript. Therefore, I invite you to respond to the reviewer(s)' comments and revise your manuscript.

Please also upload both clean and tracked changes versions of the manuscript.

To start your revision now, click the link below:

*** PLEASE NOTE: This is a two-step process. After clicking on the link, you will be directed to a webpage to confirm. ***

https://mc.manuscriptcentral.com/prisms-ext?URL_MASK=acc08ae6332f43808f30c797de82603b

Alternatively, you may log into your author centre at https://mc.manuscriptcentral.com/prisms-ext, where you will find your manuscript under “Manuscripts Awaiting Revision”. Upon submission of the revised version, your manuscript number will be appended to denote a revision. When submitting your revised manuscript, you will be able to respond to the comments made by the reviewers in the space provided. Please use this space to document any changes you make to the original manuscript. In order to expedite the processing of the revised manuscript, please be as specific as possible in your response to the reviewers.

Because we are trying to facilitate timely publication of manuscripts submitted to Cambridge Prisms: Extinction, your revised manuscript should be uploaded as soon as possible. We expect to receive your revision by 03-Nov-2025. If it is not possible for you to submit your revision by this date, please contact the Editorial Office to rearrange the due date. Otherwise, we may have to consider your paper as a new submission.

Once again, thank you for submitting your manuscript to Cambridge Prisms: Extinction and we look forward to receiving your revision.

Sincerely,

Prof. Barry Brook & Assoc. Prof. John Alroy

Editors-in-Chief, Cambridge Prisms: Extinction

13-Oct-2025

Please also ensure your manuscript complies with the following formatting points (you can find the authors instructions on this page: https://www.cambridge.org/core/journals/cambridge-prisms-extinction/information/author-instructions/preparing-your-materials):

- Please include an Impact Statement below the abstract (max. 300 words). This must not be a repetition of the abstract but a plain worded summary of the wider impact of the article.

Done.

- Submission of graphical abstracts is encouraged for all articles to help promote their impact online. A Graphical Abstract is a single image that summarises the main findings of a paper, allowing readers to quickly gain an overview and understanding of your work. Ideally, the graphical abstract should be created independently of the figures already in the paper, but it could include a (simplified version of) an existing figure or a combination thereof. If you do not wish to include a graphical abstract please let me know.

If possible, I prefer not to include a graphical abstract.

- Please ensure references are correctly formatted. In text citations should follow the author and year style. When an article cited has three or more authors the style ‘Smith et al. 2013’ should be used on all occasions. At the end of the article, references should first be listed alphabetically, with a full title of each article, and the first and last pages. Journal titles should be given in full.

References were formatted following the Cambridge A style in Zotero.

- Statements of the following are required at the end of all articles: ‘Author Contribution Statement Financial Support’, ‘Conflict of Interest Statement’, ‘Ethics statement’ (if appropriate), ‘Data Availability Statement’. Please see the author guidelines for further information.

Author Contribution Statement: I am the only author of the article and designed and wrote the whole study. AI-assisted technologies (chatGPT) were used only to improve the English.

Financial Support: There was no financial support.

Conflict of Interest Statement: The author has no conflict of interest to declare.

Ethics statement: NA

Data Availability Statement: NA

- Please submit figures as separate files in jpg/tiff format and please ensure all main text files are submitted in an editable electronic format (word/tex).

There are no figures.

Handling Editor’s Comments to Author:

Handling Editor: Surovell, Todd

Comments to the Author:

Both reviewers raised concerns with the manuscript. Reviewer 1 finds the two parts of the paper contradictory and suggests they be better integrated, while Reviewer 2 notes that you provide limited evidence to support either position. Please pay careful attention to Reviewer 1’s suggestions for reframing the manuscript to resolve this contradiction. To address Reviewer 2’s commentary, the argument would be strengthened by incorporating additional empirical support.

I appreciate the reviewers’ comments, even when I may disagree. The revision should be clearer especially in terms of better connecting the two main parts. That should also help to see that there is a quite a bit of empirical evidence, with most references being peer-reviewed published scientific studies. Yet, I acknowledge that there are philosophical aspects, as the paper is a perspective at the boundary between science and humanities, which is why I submitted it to a multidisciplinary journal with a broad readership.

Concerning reviewer 1, there is no real contradiction: the interaction between gene and environment, which for humans includes culture, determines our behaviour today as in the past. However, we (at least those of us with a background in biology) are now well aware of this. Awareness might lead to positive change and avert the worst outcomes of the global environmental crisis.

I am not arguing that moving from awareness to radical change is easy: it is not, and I am not optimistic that it might happen. Nonetheless, we are unlikely to move forward by denying that we, as a species whose most unusual evolutionary trait is exponentially cumulative culture, may be prone to overexploitation and destructiveness. If that is true, it does not imply genetic determinism. It might simply be that, being able to build culture cumulatively, the process becomes self-reinforcing like a positive feedback going out of control until the system breaks (the endpoint we must avoid).

Reviewer 2 raises a valid point (about ‘philosophical considerations’), but one that, in my view, only concerns the second aspect I mentioned above: whether genes and culture can interact in a positive direction in relation to the global environmental crisis.

The core of the paper begins with the assumption of a decisive human role in the Late Quaternary extinctions (LQEs) to argue that, even if human destructiveness were not an emergent property of the agro-industrial revolution, a longer evolutionary history of such destructiveness would not necessarily imply inevitability. This implication rests on a still-prevalent but misleading interpretation of genetic determinism, one that overlooks the fact that every phenotype results from complex gene-environment interactions. This is a scientific fact backed by countless empirical studies. Authoritarian religions and hyper-capitalism are part of the environment, as it is an increasingly widespread addiction to luxury living. These are aspects that we can change, but, as I said above, we might not be able to do it as quickly as needed.

Reviewer(s)' Comments to Author:

Reviewer: 1

Comments to the Author

From the abstract, the manuscript appears to aim to challenge the notion that humans are inherently destructive and instead emphasize the capacity of humans to bring about positive change. However, the manuscript’s structure and argumentation do not clearly follow this path. Instead, the first half of the manuscript focuses on Late Quaternary extinctions (LQE) as evidence of human destructiveness, eventually taking the stance that this trait is a given.

I suspect that the reviewer mistook the Impact Statement for the Abstract. The Impact Statement, which I now revised, may have focused too narrowly on the essentialist view of human nature, without better connecting it with the presumed role of humans in LQE. The abstract, however, faithfully mirrors the structure and content of the main manuscript.

That humans are destructive by nature is not a given: it is a position common in the sociological interpretation of the implication of a decisive role of humans in LQE. My stated aim is to demonstrate that, even if that is the case and, therefore, human destructiveness is not a feature of modernity, this “trait” is part of an extended phenotype that cannot be simply and merely determined by genetics. Thus, destructiveness can be a feature of our evolutionary history and yet is neither deterministic nor inevitable.

The manuscript then proceeds to describe modern environmental impacts associated with human actions, identifying mass consumption and exponential population growth as significant factors contributing to the current ecological situation. The conclusion appears to be that human “culture” offers a means to address the environmental degradation that is occurring today. However, the consequence of this argument structure is that the link between the supposed human-caused LQE extinctions and modern environmental degradation breaks down. This leaves the manuscript feeling like it is telling two separate stories.

I would put it differently, which is my main point in the article: it’s the interaction between gene and the environment, which for humans crucially includes culture, that can be either negative or positive for the planetary ecosystem. The outcome of this interaction has mostly been destructive, to the point of us being the only species able to set life on the path of a mass extinction. The question is: can we change, stop degradation and start living a more ecologically balanced life?

As with the reviewer, it seems that for many scholars the answer is positive only if human impacts started with domestication and agriculture (later magnified by the industrial revolution), and negative if those began in ancient hunter-gatherers societies. Why? Because that would indicate that, since our early history as a species, we have never lived in balanced with the ecosystem.

Yet, if hunter-gatherers truly were the decisive factor in the LQE of much of the megafauna, weren’t they so impactful because of their socially transmitted technological achievements? Then, the root of these impacts is again cultural evolution and, by definition, culture is not a genetically inherited trait. Culture made us ecological dominant, but luckily culture changes faster than genes.

That our impacts have been largely negative does not bode well for the future; yet it cannot be taken as evidence of a genetically determined destructiveness, as suggested by essentialist interpretations. For most of our evolutionary history, we were unaware of the full extent of our ecological impact. Now we are, and that awareness itself is a cultural development. Rather than offering new means to exploit natural resources, this cultural shift provides the knowledge needed to change course, adopting a long-term perspective that places ecosystem health at its center. Whether this awareness will be enough to trigger the transition required to avert the crisis, and whether such a transition will occur in time, are different questions. I am not optimistic, but neither am I deterministic.

I tried to stress even more the link between the two parts. Changes are minor, as suggested by the editors, but they should make more evident the connection of the non-deterministic implications of potentially human induced LQE.

The conceptual fingerprints of the overkill model are found throughout the manuscript. Unfortunately, the use of overkill in the argument automatically forces the discussion of human impacts to destructiveness as an inherent and fixed trait.

The reviewer seems to fall in the trap of genetic determinism.

For instance, as I wrote in the article, organisms evolve to maximize fitness. No doubts the genetic predisposition for this is strong in us too (otherwise, probably, we would not be here). Yet, in the current highly modified environmental context, one which is largely driven by culture, fertility rates are decreasing almost everywhere and are negative in several western countries. This is not happening because of coercive measures. In fact, in some of these countries, my own included, there are incentives to have more children. Yet, for a complex variety of reasons, most couples decide to have fewer children. An inherent propensity to fertility does not prevent its reduction: culture overcomes genetic predisposition.

Can we achieve an analogous change in the (cultural) environment, so that our ecological impacts are rapidly reduced and brought under control? In theory, this is possible, and this possibility is not negated even if anthropogenic impacts may have accompanied most of our evolutionary history.

However, the second half of the manuscript argues for the ability of humans, through culture, to overcome environmental challenges. In other words, the first half is based on the argument of the fixed nature of human behavior, while the second half leans into the idea that human behavior can strive for solutions. To be logically consistent, the manuscript could retain the LQE premise with its fixed human behavior and dedicate the second half to documenting modern destruction, with no solutions or hope.

A predisposition for a certain behaviour does not mean that the behaviour is “fixed”. See my example above about fertility and reproduction.

As I wrote in the paper, “We may be ‘destructive by nature’ with a long chain of anthropogenic extinctions that begins 50,000 ya, if not earlier. However, ... the most profound changes for us and the planet are much more recent and, mostly driven by culture”. If destructiveness is a ‘by-product’ of a predisposition to cumulative culture, the predisposition remains but the direction of the cultural trajectory can be changed. Will that happen? Highly unlikely does not mean impossible.

Alternatively, the manuscript could be reoriented to shift the prehistoric narrative to focus on human behavior as situationally diverse and creative to support the potential for humans to mitigate environmental conditions.

Human behaviour varies widely, I agree. Also the responsibility for the current crisis varies, as I acknowledged in an entire section (“Change and responsibility: Who bears the burden for the 21st-century environmental crisis?”) dedicated to this topic. Yet, the terrestrial megafauna collapsed as it never happened before in the entire Cenozoic and today ecosystems, devoid of a large and representative megafauna component, are further massively degraded.

Of course, we cannot exclude that LQE were exclusively driven by natural climate change. That seems increasingly unlikely. Regardless, I stated why I make this assumption and fully acknowledged its uncertainty.

For clarity, I added a sentence in the last section in which I stress again the diversity of human societies but also the pervasive evidence of ancient impacts across all terrestrial habitats.

Much of what is described in the second section on the impacts of LQE would have occurred whether it was caused by climate change or human predation.

This is true, but the scale and ecological consequences of the event must be made clear: it was not merely the disappearance of a few hundred large animal species; the entire terrestrial ecosystem was fundamentally transformed by their extinction. For me, this was not a trivial realization. This section is also fully referenced and is not ‘philosophy for its own sake’, but an argument grounded in evidence.

The quotes and examples referring to indigenous peoples should be removed (page 3, line 43; page 8, line 40; page 11, line 14). The first paragraph suggests that the concept of “survival of the fittest” justifies destructive colonial actions. Given the positionality of Western research relative to indigenous peoples in colonial contexts, we should be thoughtful about using their words and experiences in our argumentation.

As in Ghosh (2021), from whom I borrowed that quotation, its purpose is precisely the opposite: to illustrate that technological superiority in warfare may be a factual condition, yet it provides no moral justification for domination, colonialism and genocide. By the same reasoning, humanity’s technological dominance over “non-human nature” offers no justification for environmental destruction or ecocide.

I regret that the reviewer overlooked this conclusion, which I explicitly emphasized multiple times, as I also stressed the role of inequality and the overwhelming responsibility of Western societies. For instance, among many other relevant quotes from the paper:

• “Today, few would accept genocide, and ecocide, as inevitability of being humans”.

• “The demise of native populations conquered by a technologically superior invader is just a special within-species case of a broader phenomenon of human ecological dominance”.

• “Palaeolithic hunter-gatherers were initially unaware of the consequences of their impact. Unlike them, in sharp contrast, but similarly to General Carleton, we know we are leading other life forms to extinctions”.

• “If anything, clarifying similarities and differences between human-driven extinction events, and the related large-scale loss of ecosystemic balance, contributes to understand how the massive inequality within and among nations in the more recent history of humanity is at the base of the current environmental emergency and, thus, a priority to address for any effective solution to the planetary crisis”.

• “Furthermore, the benefits of ‘progress’ were and are highly unequally distributed, and originate from colonialism and economic imperialism (Ghosh 2021; Hickel et al. 2022; Lewis and Maslin 2020; Ross 2017) that can be seen as an intraspecific variant of supremacism … Environmental impact interacts with social injustice, as it hits harder the poorest among people and nations, ironically, precisely those who contributed the least to the crisis (Chancel 2022; Fletcher et al. 2024; Wilkinson and Pickett 2024)”.

There is also an entire paragraph, where I cite Segerstråle, explaining why inferring moral judgement from scientific knowledge (including evolutionary theory) is often likely to mislead.

Thus, the parallel I draw is essential to the argument that supremacist views are always morally wrong, whether they occur within our species (as in gender, racial, or religious hierarchies) or across species (as in anthropocentrism, speciesism, or exceptionalism). While I left the main quotation, I have now have removed part of the sentences that were misunderstood. I hope this helps, but I recognize that deeply held preconceptions may continue to shape interpretation. This challenge applies to the paper as a whole: it is an observable fact that modern humans have become a geological force, but it does not follow that such power morally justifies our dominance over life and ecosystems.

Much of the critique of modern societies, such as “the will to dominate” (page 15, line 52) and supremacy over nature, is related to Western ideologies rather than a global trait of humans. This discussion itself could further support the argument that human beliefs and actions vary. However, as it stands, it could be used to support the fixed nature of humans.

There is an entire section making clear that it is mostly Western industrialized societies that are to be blamed for the current crisis. Sadly, the vast majority of modern nations are following the Western example.

I am not arguing that hunter-gatherers, if they really caused the collapse of the LQ megafauna, did it with a “will to dominate”. If that happened, there was neither intentionality nor (at least for most of the event) awareness, as I explicitly wrote. Nonetheless, human impacts are so profound, widespread, and historically deep that it is now widely acknowledged to be difficult, if not impossible, to identify any truly pristine terrestrial habitat (see added sentence and relative references). Even the Amazon is revealing extensive evidence of past anthropogenic influence. The human footprint on the environment appears virtually everywhere. Yet, as I noted, agriculture first and the Industrial Revolution later amplified these impacts to an unprecedented degree, with a quantitatively demonstrated predominance of Western nations in driving this transformation.

There is an argument that group-level selection may occur within humans when we think of solutions related to mitigating the tragedy of the commons. Laws, policies, and cultural norms become a means of constraining individual behavior to benefit the group. Thus, it could support a way for humans to reduce their environmental impact and become more a part of nature. One thing that humans do that stands out is that we excel at creating a great deal of intra-generational phenotypic variability for evolutionary processes to play around with. To constrain our views on human potential as fixed and hard-wired seems like a limiting belief we should avoid.

I agree on the conclusions in the last two sentences. That is indeed the point I make when I argue that there is a potential for positive change.

I prefer to avoid referring to group-level selection. I am not an expert. The concept remains debated and, at least for me, difficult to grasp in all its theoretical detail. Moreover, I am not sure that group selection could effectively be applied to global cooperation. By definition, it operates through competition among groups and therefore tends to promote in-group cohesion rather than inclusiveness and diversity. Even if group selection did play a role during human evolution, it would have acted in small, relatively cohesive societies. It is therefore not straightforward to see how such dynamics could apply to large, complex, and institutionally rigid nation-states. Yet, as I wrote, I am not an expert, may be wrong but prefer to avoid a digression on such a complex issue.

Reviewer: 2

Comments to the Author

This sort of philosophical musing about the long-term meaning or implications of events and processes in the past does not resolve issues raised in the text. There are no empirical facts, no testing of hypotheses, no falsifying of hypotheses – and therefore no scientific support for either of the polarized positions implied in the manuscript – “anthropomorphic supremacy” (evolution of inevitable human destructiveness) or something else (to be aspired to, such as changing human views towards restoring/conserving ecosystems). Authoritarian religions and hyper-capitalism direct humans to exploit their supremacy, which prehistoric Homo sapiens also might have been in the process of accepting, especially when globally expanding their range.

Anthropogenic dominance is a fact. When did it start? The paper acknowledges that we do not yet know if it is a feature of modernity (since the agricultural revolution), whose negative consequences accelerated with capitalism and industrialization, or something with a longer history. My point is that, even if it had a longer history (which we cannot exclude – evidence provided in the article, including about its large uncertainties), that does not imply inevitability. Why? Because our behaviour is phenotype; phenotype is product of the interaction of genes and the environment, including culture which creates our hugely extended phenotype (evidence provided here too); and, therefore, assuming we find a way to do it (the hardest part), we have the potential to change our behaviour, make it less destructive and become part of a balanced ecosystem.

I agree with the reviewer: this is a rather philosophical paper; it is not a test of hypotheses. Whether this is compatible with the journal aims, it is an editorial decision, positive or negative, that I can only accept.

Recommendation: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R1/PR7

Comments

Thank you for addressing the reviews in your revised manuscript. I am happy to accept the manuscript for publication.

Decision: Destructive by nature? What human-driven extinctions of mammoths and mastodons mean for today’s planetary environmental crisis — R1/PR8

Comments

No accompanying comment.