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Animal cultures: A triumph or pitfall of naturalism?

  • Filip Jaroš
Published/Copyright: June 20, 2025

The central concern of this article is the phenomenon of so-called animal cultures, which are proving to be a critically sensitive point in the problem of the relationship of nature and culture. The discovery of cultural diversification in the behavioral manifestations of apes and other higher animals is considered by evolutionary biologists as a triumph of naturalism: biological science has shown that humans and their culture do not represent an anomaly, as tradition, in the form of handing down skills and customs, also exists among non-human creatures. Empirical findings on the variability of traditions among individual groups of chimpanzees – the species genetically and behaviorally closest to humans – accord well with Darwin’s conception of evolution as incremental changes that can be seen in both the physical and mental aspect. At the same time, however, doubts of a theoretical and philosophical nature rise up, exposing the pitfalls of naturalism. Is it admissible to speak of a mental aspect in “full-blooded” naturalism? Is not the concept of animal culture itself somewhat dubious, if naturalism is understood as a transfer of phenomena to the level of physical processes? How does the agenda of naturalism account for the fact that studying the diversity of traditions used by chimpanzee societies in inhabiting the natural world requires an ethnographic approach? Doesn’t the use of methods from the domain of the social sciences in the study of chimpanzee societies suggest that this animal has achieved a certain degree of freedom as opposed to natural necessity?

In this article, I assemble arguments to show that animal behavior cannot accurately be described by the means of a physicalistic biology. I do this in two steps: first, I call attention to the indeterminism of individual behavior, which in modern theoretical biology is usually included under the more general concept of agency. I then observe that the behavior of many so-called higher animals includes a social component that reflects both the variability of behavioral manifestations among different groups of the same species and their uneven distribution within a given community. In recent decades, the sophisticated social forms and relationships in a range of vertebrate species have been designated as animal cultures and have become the subject of intensive study (Laland and Galef 2009).

The impossibility of describing animal behavior in terms of physical systems is related to the view of the organism as having a dialectical relationship with its surroundings and its development as being shaped by a full range of epigenetic processes (Švorcová and Markoš 2019, Švorcová 2023). Proponents of the extended evolutionary synthesis emphasize that organisms are not passive objects of selection processes, but agents who themselves create selective environments. In the communities of certain higher animals, we can speak of collectively-formed cultural niches, created by a range of behavioral manifestations, through which they inhabit and alter their surroundings (e. g., chimpanzee groups who pass down skills for cracking nuts that grow in their area). In this article, I will defend the belief that this theoretical expansion of the Darwinian modern synthesis represents a suitable framework for philosophical anthropology, which aims to reexamine the position of humans in relation to other organisms.

In the following reflections, I take inspiration from the approach of J. Fischer, who sees philosophical anthropology as an independent school of thought founded by German-speaking researchers in the first half of the twentieth century, particularly Scheler, Plessner, Gehlen, Rothacker, and Portmann (Fischer 2009, p. 154; Fischer 2022, p. 22). The common thread of these approaches is a dual view of man, who is understood as a natural being and cultural being at the same time. This dual designation gives rise to the special position of humans in nature (die Sonderstellung des Menschen). Plessner and Portmann were aware that humans, seen as part of nature, are understandable through the means of biology, yet this comes with the danger that a fully naturalistic approach will be applied to the domain of culture as well. Both therefore tried to outline a form of “philosophical biology” that would be open to phenomena associated with the humanities (e. g., expressivity and aesthetic phenomena; cf. Grene 1968). It is precisely as a complement to this type of biology that philosophical anthropology can develop – one that will not place humans in direct opposition to nature but will approach them as a specific part of nature that requires new levels of explanation (cf. Fischer 2005).

Fischer (2014) characterizes philosophical anthropology as a paradigm that seeks a third way between naturalism and culturalism, where naturalism in biology is understood as the application of Darwinian selection theory not only to non-human organisms but to humans as well. Portmann and Plessner were also critical of the tendency toward mechanism and reductionism that was characteristic of mainstream neo-Darwinian thinking at the time. I will demonstrate, however, that Darwin’s biological thought can be reasonably interpreted in a non-physicalistic way: animals are treated in his work as intentional agents, which is generally in agreement with the approach of Continental biological thinkers in the first half of the twentieth century (H. Jonas, J. von Uexküll, F. J. J. Buytendijk), who directly attributed a space of interiority to animals. More significantly, the reciprocity of an organism and its environment that is manifested in developmental processes has become a postulate of the extended evolutionary synthesis, where Darwinian thinking is gradually blunting the edges that irritated philosophical anthropologists in the era of the Modern Synthesis.

Whether the dynamics of human culture can satisfactorily be explained in terms of Darwinian mechanisms is a topic of increasing debate.[1] In his article “Culture extends the scope of evolutionary biology in the great apes” (2017), Whiten further extends this question to animal cultures. A space is opening up to reexamine the assumptions of current cultural primatology through philosophical inquiry. It is no longer merely a question of whether naturalism, given the complexity of human culture and the inexhaustibility of action in a free individual, is a suitable framework for anthropology. We are faced with the fact that the behavior of great apes (and other animals) in collectively formed cultural niches transcends “natural necessity” – naturalism is in need of philosophical reexamination within biology, which can no longer be understood as a purely natural science.

The question of whether animals can be understood purely from the domain of nature has already been raised by Burgat (2006). I share her conviction on the autonomy of animal behavior and, like her, am willing to speak in this connection of various degrees of freedom. I also share her belief that the founders of philosophical anthropology (including F. J. J. Buytendijk) underestimated the abilities and social life of animals. Burgat, however, does not refer to the phenomenon of animal cultures – in ascribing freedom to the behavior of animals as well as humans, she seems generally motivated by an ethical conviction that the value of human and animal life is the same, rather than by biological knowledge. This ultimately leads her to avoid naming any difference between humans and non-human animals. My article, on the other hand, assumes the existence of an anthropological difference – i. e., an array of properties or relations that create the particularity of human life and culture.[2] In the spirit of the duality between philosophical biology and philosophical anthropology, I will here focus mainly on the former, particularly on defining a common theoretical framework for animal and human ontogeny. Only against a background that recognizes parallels between human and animal cultures will it be possible to meaningfully rethink the problem of anthropological difference.

Since animal cultures demonstrate a range of characteristics that traditionally have been attributed only to human culture, it is necessary to think more deeply about how the concept of culture has been constructed. The background for the differentiation of positions is the poststructuralist anthropology of P. Descola, which shows that the division of nature and culture is taken to be self-evident only in the modern Western culture. Descola (2013) convincingly demonstrates that this dichotomy is associated with the belief that non-human beings (objects, entities) are not endowed with interiority. This is closely related to the conception of human culture as a unique domain that has no parallel in the animal kingdom. In section 1, I give a precise definition to these two pillars of Western ontology, which Descola calls naturalistic.

The dichotomy that leads to a strict division of the sciences and the humanities has been seen as a problem that could be overcome by programs striving for methodological unity, whether in the form of naturalistic or culturalistic monism. In the course of the 19th and 20th centuries, Darwinism proved the most influential program, striving for a naturalistically founded monism which, in order to be complete, however, would need an explanation of human interiority. I intend to show that Darwin himself did not seek to deny humanism, which is grounded in the existence of human interiority (especially in the area of morality) but also that his evolutionary approach could not get by without methods that recognize interiority in the behavior of higher animals as well (section 2).

Investigation of the space of animal interiority is developing dynamically in connection with observation of various animal cultures. While most naturalistically-oriented philosophers understand Darwinism through a monistic prism, current research into animal cultures takes place against a background of various theoretical approaches, which all, nonetheless, assume a strong role for cultural inheritance. This is usually understood as a counterpart that interacts with genetic inheritance (dual inheritance theory, Richerson and Boyd 2005), though epigenetic approaches that distinguish multiple levels of inheritance (extended evolutionary synthesis, Jablonka and Lamb 2014) are increasingly gaining ground. In any case, the existence of cultural inheritance in animal societies is a factor that casts doubt on human cultural exceptionalism: it is apparent that non-human societies are active creators of the world, which they structure and inhabit in many different ways (on cultural diversity, see section 3).

A recognition of the interiority of animals on one hand and human cultural exceptionalism on the other is a paradigm of the discipline that Fischer (2022) calls philosophical anthropology. In section 4, I introduce the classic approach of philosophical anthropology, famous for its sensitive and empirically-grounded description of animal interiority and its critical comparison of the ontogeny of great apes and humans. A. Portmann recognizes in higher animals an ability to transmit traditions and customs, but he posits a clear distinction between the cultural production of humans and the unconsciously created artifacts of other animals (e. g., birds’ nests). In the conclusion, I reflect critically on what the phenomenon of animal cultures and the theoretical understanding of them in the extended evolutionary synthesis means not only for Portmann’s anthropology but also other programs of cultural exceptionalism based on ontogeny and phylogeny (Tomasello).

1 The Western dichotomy of nature and culture: Naturalism and humanism

Until recently, the intellectual climate of the West has been shaped by a firm belief in the dualism of nature and culture. The line of demarcation lies between the nature of animals “as the place of instincts, impulses and ‘necessity’” and the culture of humans “as the domain of rationality, language, choice and freedom” (Michelini 2020, p. 123). Descartes’ conviction that man is the only being which, through its own characteristics, has surpassed natural processes became formative for modern science: the world of matter demonstrates regularities that can be described by natural laws, whereas the human mind creates its own immaterial world. At least since the 18th century, human knowledge has been divided in two directions with very different methodologies: physics, supported by mathematics, became the model for the natural sciences, while the humanities relied primarily on records of human thought and action, making comparisons and determining genealogies.

The Frech poststructuralist anthropologist Descola considers a belief in the sharp duality of nature and culture to be a defining feature of Western ontology, which he terms naturalism – to distinguish this from other versions of naturalism, I will consistently refer to this stance as naturalistic ontology. In accordance with the general designation of naturalism, Descola’s naturalistic ontology is based on the conviction that the world is formed according to universal physical laws. Descola (2013, p. 182) further attributes to naturalistic ontology a belief in the unique position of human culture: cultural specifics are the products of intrinsic capacities unique to humans. Culture and nature are each autonomous, disjunctive domains whose demarcations are mutually dependent.

Descola’s characterization of naturalistic ontology is based on his belief that the human mind, everywhere in the world, assesses surrounding objects according to a dualism of interiority and physicality, which he defines thus:

… ‘interiority’ refers to a range of properties recognized by all human beings and partially covers what we generally call the mind, the soul, or consciousness: intentionality, subjectivity, reflexivity, feelings, and the ability to express oneself and to dream… . Physicality, in contrast, concerns external form, substance, the physiological, perceptive and sensorimotor processes, even a being’s constitution and way of acting in the world, insofar as these reflect the influence brought to bear on behavior patterns and a habitus by corporeal humors, diets, anatomical characteristics, and particular modes of reproduction. (Descola 2013, p. 116)

Different ontologies can be categorized according to whether they assess the interiority or physicality of surrounding objects through the prism of similarity or difference. For example, among the Amazonian Native American ethnic groups there is a prevalent belief that every object has an inner aspect. The diversity of things comes from their physical aspect – the “bodily garment” that clothes them. Descola calls this ontology, which attributes life to everything from mountains to human artifacts, animism. According to him, Western naturalism uses an orthogonal classification. All objects can be uniformly described in physical terms, whether it be ne natural objects, animals, people, or human artifacts. Modern science typically believes all material objects to be invariably subject to mechanical laws. The heterogeneity of objects unfolds strictly according to the differentiated evaluation of their interiority. For Descola, it is typical of naturalistic ontology to attribute interiority only to humans. Cartesian dualism fits well into this picture, as it sees not only inanimate natural objects and human artifacts, but also animals as extended things. Man alone is endowed with interiority in the form of a soul, which in the Christian context is believed to be immortal.[3]

In Descola’s view, naturalistic ontology is dualistic: the world of natural laws does not apply to the domain of the human mind, and therefore the principles of humanism can be cultivated in the realm of values. Traditional humanism arises from the belief that the sphere of human action is an expression of free will, as opposed to natural processes, whose dynamics are independent of humans and which therefore cannot sensibly be subject to moral judgments. Physicalism in the domain of nature and freedom in the domain of culture are reflexive positions that are not mutually exclusive. Although the sharp dichotomy of nature and culture is perceived as a philosophical problem, it is routinely accepted in the realm of ethics and in judicial practice.

In sum, Descola characterizes naturalistic ontology as resting on two pillars. Pillar 1 (physicalism): non-human creatures, natural objects, and human artifacts are not endowed with any sort of interiority. Pillar 2 (cultural exceptionalism): humans are the only species that engenders cultural differences between groups. In the following sections, I will consider to what extent a) Darwin’s conception of animal behavior and b) contemporary Darwinism correspond to this characterization of naturalistic ontology in relation to the phenomenon of animal cultures. It turns out, surprisingly, that Darwinism, generally regarded as the most successful naturalistic approach in the West, has helped to undermine both pillars of naturalistic ontology through its treatment of these issues.

2 Darwin and physicalism

To examine the validity of the first pillar of naturalistic ontology in relation to Darwin’s work, we should first ask whether Darwin attributes animals with interiority. The answer depends on how we interpret Descola’s assessment of interiority in relation to Darwin’s account of animals and humans, which also relates to the question of whether we assess Darwin’s epistemology as dualistic or monistic. I submit that an understanding of interiority as the domain of mind, consciousness, and emotion, together with a phenomenal account of animal behavior, will lead us to an affirmative answer: it presupposes the duality of phenomenal description and scientific theory. If, however, we understand Darwin’s theoretical thinking on the basis of monism, the duality of physicality and interiority loses its original meaning, and a monistic picture of the world leads not only to a rejection of interiority in animals but can end up casting doubt on the freedom of behavior in humans. I will hereafter refer to this coherent, naturalistically (or materialistically) informed monism as physicalism.[4]

Descola (2013, p. 199) gives preference to the physicalist interpretation when he judges Darwin’s Descent of Man (1871) to be a canonical version of the agenda of incorporating culture into nature.[5] In accordance with a materialistically-monistic worldview, interiority here no longer has the validity of an independent explanatory principle and can be converted to physicality in the form of mechanical laws. Processes taking place in human culture therefore do not qualitatively differ from processes in the natural world – they can be subordinated to the universal laws of nature, and culture can thus be incorporated into nature. Descola here mentions the principle of natural selection, which Darwin applied both to organic variation and social instincts.

There are further reasons to evaluate Darwin’s program through the lens of physicalism. In the first edition of The Origin of Species, Darwin cites Whewell’s motto on the need for scientific description of the material world to establish general laws (Darwin 1859). Many other instances show that Darwin took physics to be the model for natural science. Natural selection is presented in The Origin of Species as a physical process with universal validity, directing the physical form of organisms as well as their mental constitution and social structures. As for his theoretical view on the nature of mental processes, Darwin believes them to have a physical origin – he suggests, for example, that aesthetic taste can be derived from the physiological properties of the nervous system (Darwin 1871, p. 333).

To this assessment it must be added that in his thinking about humans, Darwin in many ways preserved the agenda of humanism (in Descola’s original assessment of naturalistic ontology, the world of natural laws applies to non-human animals but not humans). Darwin was an ardent supporter of the abolition of slavery, as he believed all humans are equally born free. In general, he held predominantly liberal views on moral issues, was an optimist when it came to the development of the human race, and was an enthusiastic proponent of the idea of moral progress in history. [Here and elsewhere, his values are close to Unitarianism (see Reed 2011)]. His reflections did not touch on the space of the human soul – he was far from seeing humans as sophisticated machines lacking free will. Those who accused Darwin of reducing humans to mere apes were mistaken: I will incrementally show that the outcomes of Darwin’s work signify the elevation of animals towards humans, rather than the “bestialization” of the human race.

The following reflections are dominated by my conviction that Darwin’s primary intellectual legacy – the evolutionary link between humans and animals – is grounded not in a programmatic physicalism but in a dualistic approach to living organisms. It is useful to recall that Darwin, rather than a scientific theorist, was a brilliant expert on animal behavior. One sees this when reading The Expression of the Emotions in Man and Animals (1872), the foundational work in comparative psychology and ethology, which still today remains a testament to the great observational talent of the British naturalist. The attribution of a mental life to animals was an important condition for the vision of evolution without supernatural intervention: continuity on the corporeal level only (e. g., the similar anatomy of humans and apes) would leave open the question of the origin of the human mind (cf. Jaroš 2017). Moreover, a direct treatment of what Descola calls interiority is to be found in The Descent of Man:

… man and the higher animals, especially the Primates, have some few instincts in common. All have the same senses, intuitions and sensations – similar passions, affections, and emotions, even the more complex ones; they feel wonder and curiosity; they possess the same faculties of imitation, attention, memory, imagination, and reason, though in very different degrees. (Darwin 1871, p. 48 – 49)

Ethology and comparative psychology necessarily deal with intentionality, feelings, and the ability to express oneself, all of which belongs to Descola’s conception of interiority. In the field, ethologists must always rely primarily on their senses and experience in order to grasp the meaning of the events they are witnessing. Significantly, interiority and physicality (which includes morphology, for example) are interconnected; thus, we understand the emotions of animals just as we do with humans – primarily through expressions and body language. An ethological theory can employ a model that is physicalist (natural selection) or cybernetic (fixed action pattern; Lorenz 1981), but when it is put into action, the scientist must be able to understand phenomena that belong to the preserve of interiority.

Darwin shows himself to be a great thinker of animal interiority. Above all, the “higher animals” appear in his work as true agents: “Animals may constantly be seen to pause, deliberate, and resolve” (Darwin 1871, p. 46). Even if he later decided to downgrade this phenomenal level by portraying it as only an epiphenomenon of physical or neurochemical processes, in terms of his evolutionary vision, he would have to make an analogous reduction in the case of human mental processes and perhaps human culture. In any case, Darwin’s treatment of interiority implies a twofold process: firstly, an elevation of interiority in animals toward human behavior, and secondly, a “biologization” of the human mind, which had until then been understood to be voluntaristic.

To summarize thus far: I maintain that the achievements of physicalism in mathematically grasping and describing natural processes by no means extend to the entire realm of nature (physis) but only to the inanimate world. Organisms require a separate approach, as closer experience of them reveals a multitude of characteristics that correspond to Descola’s conception of interiority. The “certainties of naturalism” (Descola 2013, p. 172) are called into question the moment biologists decide to study animal behavior in the field, which requires a completely different epistemology than laboratory research, experimental organisms can be compared to automatic machines.[6] Biology has been epistemologically emancipating itself from physics since Darwin’s time, without being fully aware of it. As a consequence, it will cease to be clear what it means to “naturalize” a given phenomenon, as there can be multiple levels of explanation in natural science (cf. Sterelny 1996). I am inclined to conclude that Darwin’s treatment of animal interiority amounts to a challenge to the first pillar of Descola’s naturalistic ontology.[7]

3 Animal cultures: Between natural necessity and human creativity

When Descola (2013, p. 72 – 78) describes the autonomy of the domain that inhabitants of the West have traditionally called culture, he relies on the concepts of ethnologists like E. B. Tylor and F. Boaz, whose definitions of culture, however, assume that cultural phenomena are found only among humans. If we want to examine the validity of the second pillar of ontological naturalism (i. e., cultural exceptionalism), we need to use a conception of culture that at least potentially allows for its occurrence in nonhumans. Descola (2013, p. 182) does, in fact, concede something of the sort: “To cut naturalism down to size, it would be necessary to show that the chimpanzees draw upon psychic resources identical to our own when they engage in cultural activities.” The problem here, of course, is the word identical: in the spirit of Descola’s structuralist approach, it would be more appropriate here to ask whether this amounts to similar or dissimilar psychic resources. Only dissimilar psychic resources in humans and chimpanzees can justify the insistence that only humans have culture in the true sense, derived from a categorically different interiority. How then to understand culture so that it is not by definition only human, and how to determine what psychic qualities chimpanzees are endowed with?

A reversal in biological research occurred in 1960 when J. Goodall observed chimpanzees making and using tools. The techniques used turned out to be traditions specific to particular chimpanzee groups, which raised the question of how such knowledge is propagated. Goodall was able to demonstrate the existence of social inheritance across chimpanzee societies – i. e., in cases other than parent–offspring transmission. She eventually realized that the study of chimpanzee societies requires not just observation from a distance, but acceptance by the group during her participant observation and other methods typical of an ethnographic approach (see Goodall 1990). By the late 1970s, the phenomenon of ape cultures was recognized by natural scientists and many anthropologists – McGrew and Tutin’s 1978 study, published in Man, a journal of social anthropology, can be considered a breakthrough in this regard. An ethological definition of culture focuses on the transmission of behaviors in a group: “a cultural behaviour is one that is transmitted repeatedly through social or observational learning to become a population-level characteristic” (Whiten et al. 1999). This approach assumes nontrivial cognitive abilities on the part of individuals who pass cultural practices to each other, but it holds back on statements about the nature of these abilities, maintaining that the mind is an impenetrable black box.

Today, researchers are looking at animal cultures not only in primates but elephants, cetaceans, and birds – cultural behavior is presumed in hundreds of vertebrate species (Laland and Hoppitt 2003). Vocal traditions have been documented in cetaceans (humpback whales are particularly known for this) and passerines. Cebid monkeys and chimpanzees use rocks to crack nuts, and cockatoos can prepare small pieces of wood to use for obtaining food. The grooming hand-clasp performed before mutual grooming in some chimpanzee societies (de Waal and Bonnie 2009) can also be counted as a cultural sign. As the character of this gesture has no direct physical relationship to the behavior it precedes, it amounts to an arbitrary sign that can be classified as a symbol (cf. Jaroš and Pudil 2020, p. 171).

The existence of ape and other animal cultures presents adherents of Darwinian gradualism an opportunity to examine how human culture may have come about in the first place. In this respect, animal cultures represent a triumph of naturalism: the domain of human culture appears as a special instance of social structures that can be found in higher animals – a domain accessible to natural-scientific inquiry. But the question remains of how to evaluate the psychic makeup of chimpanzees and other great apes in comparison with humans. The answer depends primarily on whether the research is conducted by ethologists (who focus directly on behavior) or comparative psychologists working in the laboratory (who focus on cognition deduced from behavior): while the former tend to look for analogies between animal and human behavior, the latter implicitly proceed with an agenda of finding anthropological difference (cf. Jaroš and Maran 2019).

Field primatologists like Boesch (2012) attribute culture to chimpanzees and other great apes because, given the many analogies (hunting in groups, frequent tool use) and the close evolutionary relationship, it is economical to assume that the minds of humans and apes are similar [de Waal (1999, p. 259) talks about the principle of evolutionary parsimony]. On the other hand, comparative psychologists concentrate on laboratory research into the cognitive processes that underlie the modes of cultural transmission. Tomasello (1999) shows that children, when using tools, copy exactly the sequence and implementations of steps demonstrated by their instructor (imitation), while chimpanzees are able to understand the intention of the demonstrator and then “willfully” employ whatever tactic leads them to this goal (emulation). Humans thus have greater accuracy and fidelity in the transfer of information and skills; this, together with the highly organized nature of their groups, results in the cumulative character of their culture (Tomasello, Savage-Rumbaugh and Kruger 1993; Boyd and Richerson 1996).

Independent of the dispute over the cognitive background of cultural transmission in apes and humans, it is useful here to reflect on the extent to which the phenomenon of animal cultures corresponds to the interpretation of naturalism in this text. I consider the conception of animal mind used in cognitive ethology and comparative psychology to be far from the image of nature as a necessary process whose course is described by the laws revealed by science. This is primarily in regard to the singularity of the moment when a new behavioral trait appears. While neo-Darwinism presupposes a random genetic mutation (or several) in the case of physiological adaptations, which are then fixed by natural selection, behavioral adaptations are not a matter only of genetics but are primarily the result of the activity of the individual. Biologists do not hesitate to speak of innovation, and it is believed that individuals who have a tendency toward exploration and playfulness are distinguished by an ability to produce innovation (cf. Morand-Ferron et al. 2011).

It is also significant that the ethological definition of culture (Whiten et al. 1999) draws on dual inheritance theory, which indicates an opposition of genetic and cultural inheritance. Culturally conditioned behavior characteristically is not an immediate (genetically fixed) reaction to events in the surrounding environment, but is observed in, and learned from, other individuals in the group, different groups producing different variants. The plasticity of animal culture in contrast to genetically determined behavior thus recalls the dualism that Descola and others apply to the dichotomy of culture and nature in the case of humans. If we understand genetic and cultural inheritance as distinct, even if not totally separable processes, on the model of dual inheritance theory, then it is reasonable to conclude that animal cultures transcend biological necessity and open the possibility for a new understanding of the concept of freedom.

An ethological definition of culture rests on a synchronic perspective comparing the behavior and skills of communities separated by space. An individual possesses behaviors that are predetermined by his genetic makeup and skills that he has learned from other individuals in his group (or perhaps he was the first to acquire them). The division of hereditary transmission into two types is complicated, however, when we consider that each skill must be acquired during individual ontogeny in relation to the environment in which the individual develops. This environment, moreover, is not just a physical space, but rather a milieu that is modified by the cumulative influence of the activities of past generations. In this modified view, culture is not in strict opposition to the natural environment, but is progressively woven into it. The developmental definition of culture focuses on the ways in which the social actions of past generations enhance or facilitate what further generations learn in a collectively structured environment (Lewens 2017, p. 5). The actor here is not the individual but the entire (culturally equipped) society. We find an apt example in the environmental engineering of beavers, which demonstrates the active inhabitation of a natural environment (niche construction), rather than just passive presence.

The developmental definition of culture and niche construction are among the important theoretical postulates of the modern movement of evolutionary thought known as the extended evolutionary synthesis. This holds that animals in societies construct niches in which they live and raise future generations – this niche is part of the natural environment but also represents an adjustment to the environment toward the needs of a given animal culture (cf. Laland and Brown 2018). A niche should be understood not just as a physical space but a way in which the surroundings are represented and elements of it are selected as bearers of meaning (Švorcová and Kleisner 2018, p. 236 – 237). From an ontogenetic perspective, it is essential that the young progressively acquire the skills and traditions of their group – i. e., that they grow up in an environment that is pre-saturated with these activities. This does not only mean that there are objects around that can be used as tools, but that the young can gradually master the relevant techniques through a combination of observing older individuals and actively experimenting on their own.[8]

A fine example of active maintenance of an animal cultural niche is the cracking and consumption of nuts by chimpanzee groups in the Taï forest (Boesch 2012). An important part of Taï chimpanzees’ diet consists of nuts of the Coula tree, which can be cracked only by the use of a wooden stick (hammer) that hits a nut positioned on a stone (anvil) at a proper angle. The technique is transmitted vertically (i. e., across generations) from a mother to her offspring, and its full mastery usually is not acquired before the age of 7. It appears that the space for mastering the technique is actively created through cooperation on the part of mothers, who leave used wooden sticks in the vicinity of the infants and support them in their initial and subsequent attempts to crack nuts. Matsuzawa et al. (2001) call such dynamics an education by master-apprenticeship, which is characterized by assistance rather than active teaching. The crucial component of the education is a stimulating learning environment characterized by an atmosphere of mutuality and tolerance. Apprentices learn by active participation, and when a required technique is not difficult, active teaching by the mother is not necessary.[9]

Rather than intervening, mothers need to develop sensitivity for the progress of their apprentices and correctly assess their skill level and react appropriately. Boesch has divided the transmission of nut-cracking ability into three phases: sharing, stimulating, and facilitating. First, mothers allow their offspring to eat around 25 % of the kernels they have secured from nuts. Infants imitate the acts of their mothers but are not able to crack the nuts due to their limited physical strength; however, they might assist by collecting them and placing them on the anvil. When they reach the age of 3 – 4, their mothers stimulate them by leaving nuts suitably positioned on the anvil, so the infant can crack them by using the hammer while the mother leaves to pick other nuts. When infants reach adolescence at the age of 5 – 8, they have generally acquired the motor-cognitive abilities to crack the nuts all by themselves. Nevertheless, they still have to learn which kinds of wooden branches can be used as hammers. At this stage, mother facilitates their semi-independent activity by leaving them with the hammer and searching for new ones as well as for nuts. The youngster in the Täi forest is provided with hammers and nuts an average of 6 times per 10 minutes in a nut-cracking session (Boesch 2012, p. 136 – 137; Jaroš and Pudil 2020, p. 170 – 171).[10]

It is useful to summarize here how the possibility of free behavior is involved in animal interiority (section 2) and the active creation of niches (section 3). The first degree of freedom relates to the fact that a species-typical relation of the individual to its environment prefigures modes of behavior but does not determine them. Animal agency should be understood as a palette of behavioral acts that are based on the needs of the animal individual and which cannot be rendered as a physicalistic combination of action and reaction.[11] The concept of freedom in animal culture supposes the mind of animals as a space where invention occurs through ways of behaving toward the physical and social environment. Each society has a different set of customs and traditions that often carry adaptive meaning but which cannot be understood as a result of natural selection – individuals themselves build the selection environment for their group.

In the past few decades, research from the field has upended the conviction that the use and improvement of tools, the intentional transfer of techniques for using them, and the formation of arbitrary signs (symbols) are exclusively human capabilities. The phenomenon of animal cultures challenges the second pillar of Descola’s ontological naturalism, which reserves cultural exceptionalism for humans. If, in agreement with the extended evolutionary synthesis, we are going to understand culture as a way in which human or animal societies inhabit the surrounding environment and actively shape it over generations (so-called niche inheritance), then we will stop associating cultural processes by definition with an anthropocentric view of cognition. This section is not meant to conclude whether human mental abilities are unique but to find a phylogenetic framework in which to sensibly formulate the question of anthropological difference. Our next task will be to connect this framework with selected approaches of philosophical anthropology.

4 Portmann’s philosophical anthropology: Animal inwardness and the special position of humans[12]

In section 2, I attempted to show that in discussing animal behavior and its social aspects, we cannot overlook the intentionality, emotionality, and spontaneous expression of it. While for naturalistically-minded Darwinists the emergence of these aspects came about as a byproduct of their program of finding an evolutionary transition between animals and humans, for some of the founders of philosophical anthropology, the subject of interiority is a fundamental platform for thinking about living organisms. I will focus here on the ideas of A. Portmann, whose zoological expertise and well-known determination of the developmental type of humans as “secondarily altricial” have made him an authoritative figure for all philosophical anthropologists.

From a theoretical perspective, Portmann’s biology can be placed within the organicist movement, which emphasizes the importance of developing a nondeterministic epistemology in order to understand living beings. J. von Uexküll and Portmann postulate that organisms are subjects that actively interpret events in their surroundings. In this connection, Portmann speaks of inwardness (not only of animals but plants also).[13] Inwardness includes the ways in which a given organism relates to the world as well as the realm of experience, which derives from fulfilling the need of self-preservation and from emotional and social motivations. We can include the phenomenon of play here, which shows that an animal’s movements are not tied to “purely biological” drives such as hunger or sexuality. Although Portmann (1964, p. 142), in agreement with Uexküll, evaluates animal Umwelten in terms of biologically programmed functional cycles, he nonetheless sees moments of freedom in some animal behavior (neue Möglichkeit des freieren Verhaltens, Portmann 1968, p. 18). Thus, the domain of nature is not fully capturable by natural laws – not because human knowledge is insufficiently advanced, but because it is not a domain of necessity.

Portmann considers the complexity of animal social life to be high. He particularly notices in primates an ability of juveniles to adopt behaviors from adult individuals and speaks of their social Umwelt (Portmann 1964, p. 104) and the existence of traditions (Portmann 1990a, p. 57, 67). The fullest extent of these considerations can be found in his 1970 Eranos lecture on a zoosemiotic comparison of human speech and animal communication (available in English translation in Portmann 1990b). Here he explicitly states that animals acquire new habits through imitation and further points out that birds and mammals acquire new habitats through a process of “acculturation.”[14]

In the spirit of the posthumanist trend in the contemporary phenomenology of animality, Burgat (2006) observes that Portmann opened the gates of freedom to animal behavior. We must realize, though, that from a different perspective, Darwin contributed more to our understanding of the possibilities of animal behavior than Portmann. In postulating an evolutionary continuity also in the realm of mental abilities and speculating on the origin of human culture via natural (and sexual) selection, Darwin prepared the ground for the undermining of the second pillar of naturalistic ontology. Portmann, on the other hand, supports the thesis of a special position of humans in nature (die Sonderstellung des Menschen), which he adopts from M. Scheler. While Scheler developed this position by a philosophical method supported by contemporary findings in biology (especially the primatological studies of W. Köhler), Portmann tries to define human specificity primarily through morphology and ontogeny:

In spite of the fact that apes are precocial in form, they develop relationships with the environment more slowly than do other precocials – ungulates and whales, for instance. However, no great ape goes through a postpartum phase of transformation in which it does not acquire species-specific posture until it is exposed to social contact. Furthermore, no baby ape experiences the slow metamorphosis to world experience as perceived by the mind as we do in that first, developmentally critical year after birth, a stage that marks a clear division of the entire growth period into three distinct parts. We say that this developmental type is a special kind of ontogeny and call it ‘secondarily altricial.’ (Portmann 1990a, description to image 5.2)

The presence of a phase of distinct postnatal transformation not found in any other primate leads Portmann to the idea that humans are “prematurely” born compared to other apes – a purely precocial form of development would require the human embryo to continue in its uterine development for a further 9 – 12 months. The rapid development during this period, which corresponds to the dynamics of embryogenesis in the womb, leads Portmann to believe that the first extra-uterine year of human life is a unique phenomenon in biological evolution.[15] According to Portmann (1990c, p. 132), three key characteristics of humans – upright posture, the beginnings of spoken language, and the foundations of rational thought – are formed during the 7 – 12 months following birth. This early dynamic phase of human ontogeny occurs in an external environment rich in various stimuli, but one which is also more dangerous than the environment of the womb. To survive, a helpless human infant needs not only the attentive care of a mother but also, given the supremely social form of the human species, the support of the entire community into which it is born. This insight led Portmann to formulate the metaphor of the “social uterus” (sozialer Uterus)[16] as the space that allows the child to grow up to become a fully-fledged member of a human group, both physically and mentally.

To the basic triad of upright posture, speech, and a rational relation to the world we must add another related human characteristic – life in societies shaped by culture. Portmann (1968, p. 30) speaks of culture as a mode of expression that goes beyond natural spontaneity. A typical example of controlled expression is human speech, which utilizes symbolic expressions passed down from generation to generation. Portmann contends that culture is essential to humans and criticizes efforts to find an evolutionary phase in which they belonged anatomically and morphologically to the species Homo sapiens yet were not endowed with a complex social and symbolic life.

Here we see skepticism of the Darwinian concept of transitional links, whose hypothetical existence corresponds to the assumption of a continuous transition between individual species. According to Portmann, human culture differs from the transmission of traditions in animal societies in that it can pass on the stories of past generations to which we can consciously relate – only in the case of humans can we speak of this kind of history (Portmann 1990a, 11).

5 In lieu of conclusion: Animal cultures as a challenge for philosophical anthropology

One goal of this article was to show how modern biological thought has enriched our conception of the animal. It is evident that in order to understand animal behavior, it is expedient to replace the physicalist interpretation with the concept of agency. The phenomenon of differentiation of social, material, and symbolic life across various communities of a given species, known as animal culture, lies on the border between the research domains of the sciences and humanities. This is reflected primarily in the realm of methodology – for example, in the field of cultural primatology, natural scientists are literally required to live among primate communities in order to identify their habits and understand their significance. It can be said with some exaggeration that the discovery of chimpanzee culture signaled the end of biology’s status as a purely natural science.[17] Many interdisciplinary fields are being established today such as multispecies ethnography and human-animal studies, which represent a mix of various approaches from natural science and the humanities. These methodological transformations reflect a deeper ontological shift in the approach to animal behavior.

Finally, let us ask what this shift in the understanding of culture means for philosophical anthropology. Of the founders of philosophical anthropology, Portmann was the one most concerned with the nature of the social world of animals (see namely Portmann 1964), so it seems useful to respond to his reflections on the position of humans in relation to animals, which were introduced in the previous section. Without questioning the validity of the overall program of seeking a special position for humans in nature, I would draw attention to some moments where I believe Portmann underestimated the analogy between great apes and humans. As primatologists learn more about emotional and social factors during the early years of ontogeny in chimpanzees and other great apes and their interconnection with the cognitive and social intelligence of adult individuals, it is clear that developmental plasticity plays an important role in our closest relatives (Bard et al. 2014). The importance of physical and psychological contact in rearing infants is a constant in all primates; Bowlby’s theory of the emotional bond between a child and its caregiver is merely a special case of general need for a close emotional connection between an infant and the individual providing care. Moreover, the assumption that the rearing of an infant is bound to the biological mother is not true even in great apes, as shown by the phenomenon of adoption in chimpanzees (Goodall 1990) and gorillas (Morrison et al. 2021). Chimpanzee infants engage in play with other members of the group, including adult males, at a very young age, and their frequently disruptive behavior is tolerated specifically thanks to a general ability to perceive them as immature individuals. In light of a certain degree of acculturation seen in young apes with human foster caretakers, it is also necessary to revise Portmann’s view that “in the well-protected, newborn ape no essentially new possibilities for behavior and movement or for means of communication arise” (Portmann 1990a, p. 57). A space is opening up in which we can also glimpse the development of young apes through the prism of a kind of “social uterus,” since their maturation, too, is not a simple fulfillment of biological dispositions and has a fundamentally social character.

It is worth emphasizing that while current deliberation in comparative psychology and cultural evolution studies takes place within a Darwinian paradigm, the specificity of human culture is rarely questioned. Most mainstream authors speak of the unique dynamics of human cultural evolution, which they designate as cumulative, yet there are indications that skills are accumulated in some chimpanzee cultures as well (Whiten 2017, p. 7794). In connection with this question, specific qualities of humans are being sought in the field of social cognition. Most influential in this regard is Tomasello’s hypothesis that the anthropological difference derives from the capacity for shared intentionality. Even though he does not discuss the relationship of his work on natural history to the foundational works of philosophical anthropology, Tomasello’s linking of the agenda of a special position of humans in nature with a nonreductive naturalism won him the Helmuth Plessner Prize (cf. Fischer 2016). According to J. Fischer and H.-P. Krüger, Tomasello’s natural history of anthropogenesis fulfills the postulate of Plessner’s philosophical anthropology, which sees humans as natural and cultural beings at the same time (natürliche Künstlichkeit, Plessner 1975, p. 338).

The strength of Tomasello’s investigation of the anthropological difference is its merging of ontogenetic and phylogenetic considerations. In the face of new knowledge about animal cultures and so-called humanized chimpanzees, however, Tomasello’s conception requires a similar kind of revision as Portmann’s views, namely his underestimation of the abilities of chimpanzees, particularly in the area of social cognition. This may be due to his chosen method of laboratory research, where the abilities of chimpanzees are compared to those of children without consideration of the differences in individual ontogeny (children grow up in an environment rich in objects and activities that are unknown to the chimpanzees being tested). Leavens (2004) convincingly demonstrates that, through enculturation, chimpanzees are able to understand and actively use hand pointing as declarative gestures. This raises doubts about Tomasello’s postulate on the Machiavellian intelligence of chimpanzees, who are able to solve complex problems but are supposedly motivated by the pursuit of individual advantage, usually a better position in the group or access to sexual partners. When we consider their formation of affective bonds and wider coalitions, however, the image of chimpanzees as egoistic manipulators with selfish inclinations seems too one-sided (cf. de Waal and Bonnie 2009). Field primatologists like J. Goodall and C. Boesch have observed chimpanzee cultures of hunting monkeys that demonstrate true cooperation and culminate in sharing of the prey. In the end, I am convinced that Tomasello’s hypothesis of shared intentionality must be theoretically reexamined by means of the extended evolutionary synthesis – social-cognitive abilities are influenced by prior ontogenetic trajectories, and the relationship of the individual to the environment has a social-cultural component.

In closing, I would like to propose a path for philosophical anthropology if it will consider moving beyond the modern dichotomy of non-human nature and (exceptionalistic) human culture. I have attempted to show that the ontogenetic-phylogenetic approach, increasingly prevalent these days in the extended evolutionary synthesis, is a suitable framework for thinking about animals and humans. This approach links the space of interiority with the environment in which the relevant society lives and, at the same time, shows that the methods by which a group modifies its environmental niche are transmitted from generation to generation. Animal cultures thus represent a type of phylogenetic memory that must continually be updated in the ontogeny of the members of a given group. Redefining culture does not have to mean abandoning the search for the anthropological difference, though its place must lie outside the traditional nature–culture dichotomy. In establishing the position of humans in evolution, we must take into account the fact that not only humans but also the “higher animals” have biological predispositions to free themselves from the “natural necessity” seemingly encoded in their species-typical genetic makeup.

I wish to thank Andrew G. Christensen for the translation into English and Jan Kapusta, Jana Švorcová, Michal Hubálek, Iva Svačinová, Márton Dornbach for their helpful comments. This publication was supported by a project of the Czech Science Foundation (GA23-05374S, Reframing Philosophical Anthropology: Searching for an Anthropological Difference Beyond the Nature/Culture Dichotomy) at the University of Hradec Kralove, Philosophical Faculty.

References

Bard, K. A., Bakeman, R., Boysen, S. T., Leavens, D. A. (2014). Emotional engagements predict and enhance social cognition in young chimpanzees. Developmental Science, 17(5), 682 – 696.10.1111/desc.12145Search in Google Scholar

Boesch, C. (2012). Wild cultures. A comparison between chimpanzee and human cultures. Cambridge: Cambridge University Press.10.1017/CBO9781139178532Search in Google Scholar

Boyd, R., Richerson, P. J. (1996). Why culture is common, but cultural evolution is rare. Proceedings of the British Academy, 88, 77 – 93.Search in Google Scholar

Brentari, C. (2018). From the hiatus model to the diffuse discontinuities: A turning point in human-animal studies. Biosemiotics, 11(10), 331 – 345.10.1007/s12304-018-9329-8Search in Google Scholar

Burgat, F. (2006). Liberté et inquiétude de la vie animale. Paris: Kimé.10.3917/kime.flore.2006.01Search in Google Scholar

Darwin, C. R. (1859). On the origin of species. London: J. Murray.Search in Google Scholar

Darwin, C. R. (1871). The descent of man, and selection in relation to sex. New York: D. Appleton.10.5962/bhl.title.24784Search in Google Scholar

Darwin, C. R. (1872). The expression of the emotions in man and animals. London: J. Murray.10.1037/10001-000Search in Google Scholar

Descola, P. (2013). Beyond nature and culture. Chicago: The University of Chicago Press.10.7208/chicago/9780226145006.001.0001Search in Google Scholar

de Waal, F. B. M. (1999). Anthropomorphism and Anthropodenial: Consistency in Our Thinking about Humans and Other Animals. Philosophical Topics, Vol. 27, No. 1, 255 – 280.10.5840/philtopics199927122Search in Google Scholar

de Waal, F. B. M., Bonnie, K. E. (2009). In tune with others: The social side of primate culture. In K. Laland, B. G. Galef (Eds.), The question of animal culture (pp. 19 – 39). Cambridge, MA: Harvard University Press.10.2307/j.ctv322v4wf.5Search in Google Scholar

Fischer, J. (2005). Biophilosophie als Kern des Theorieprogramms der Philosophischen Anthropologie. Zur Kritik des wissenschaftlichen Radikalismus. In Alexandra Manzei, Mathias Gutmann & Gerhard Gamm (eds.), Zwischen Anthropologie Und Gesellschaftstheorie: Zur Renaissance Helmuth Plessners Im Kontext der Modernen Lebenswissenschaften. Transcript Verlag. pp. 159 – 182.10.1515/9783839403198-008Search in Google Scholar

Fischer, J. (2009). Exploring the core identity through the works of Max Scheler, Helmut Plessner and Arnold Gehlen. Iris, 1, 153 – 170.Search in Google Scholar

Fischer, J. (2014). Philosophical anthropology. A third way between Darwinism and Foucaultism. In J. de Mul (Ed.), Plessner′s philosophical anthropology. Perspectives and prospects (pp. 41 – 54). Amsterdam University Press.10.1515/9789048522989-003Search in Google Scholar

Fischer, J. (2016). Michael Tomasello und Helmuth Plessner. Zwei Protagonisten der modernen Philosophischen Anthropologie. In J. Fischer, Exzentrische Positionalität. Studien zu Helmuth Plessner (pp. 319 – 342). Weilerswist: Velbrück Wissenschaft.10.5771/9783845280967-320Search in Google Scholar

Fischer, J. (2022). Philosophische Anthropologie. Eine Denkrichtung des 20. Jahrhunderts. Münich, Freiburg: Karl Alber.10.5771/9783495999899Search in Google Scholar

Goodall, J. (1990). Through a Window: 30 years observing the Gombe chimpanzees. London: Weidenfeld & Nicolson; Boston: Houghton Mifflin.Search in Google Scholar

Grene, M. (1968). Approaches to a philosophical biology. New York: Basic Books.Search in Google Scholar

Jablonka, E., Ginsburg, S. (2022). Learning and the Evolution of Conscious Agents. Biosemiotics 15, 401 – 437.10.1007/s12304-022-09501-ySearch in Google Scholar

Jablonka, E., Lamb, M. J. (2014). Evolution in Four Dimensions: Genetic, Epigenetic, Behavioral, and Symbolic Variation in the History of Life. Cambridge, MA: MIT Press.10.7551/mitpress/9689.001.0001Search in Google Scholar

Jaroš, F. (2017). Darwin, Dawkins, and de Waal: Animals Like Persons or Machines? In M. Škorić, A. Kišjuhas (Eds.), Atheism and Moral Progress (pp. 143 – 168). Novi Sad: University of Novi Sad, Faculty of Philosophy. ISBN 978-86-6065-405-4Search in Google Scholar

Jaroš, F. (2021). Portmann’s view on anthropological difference. In F. Jaroš, J. Klouda (Eds.), Adolf Portmann: A thinker of self-expressive life (pp. 119 – 142). New York: Springer.10.1007/978-3-030-67810-4_7Search in Google Scholar

Jaroš, F. (2022). Portmannova filosofická biologie a antropologie. Praha: Togga.Search in Google Scholar

Jaroš, F., Brentari, C. (2022). Organisms as subjects: Jakob von Uexküll and Adolf Portmann on the autonomy of living beings and anthropological difference. History and Philosophy of the Life Sciences 44, 36.10.1007/s40656-022-00518-7Search in Google Scholar

Jaroš, F., Maran, T. (2019). Humans on top, humans among the other animals: Narratives of anthropological difference. Biosemiotics, 12(3), 381 – 403.10.1007/s12304-019-09364-wSearch in Google Scholar

Jaroš, F., Pudil, M. (2020). Cognitive systems of human and non-human animals: At the crossroads of phenomenology, ethology and biosemiotics. Biosemiotics, 13(2), 155 – 177.10.1007/s12304-020-09387-8Search in Google Scholar

Jonas, H. (1966). The Phenomenon of Life: Toward a Philosophical Biology. New York: Harper & Row.Search in Google Scholar

Klouda, J. (2021). The “new morphology” between biology and philosophy: The hermeneutic dimension of Portmann’s thought. In F. Jaroš, J. Klouda (Eds.), Adolf Portmann: A thinker of self-expressive life (pp. 199 – 218). New York: Springer.10.1007/978-3-030-67810-4_11Search in Google Scholar

Kurismaa, A. (2021). Revisiting basal anthropology: A developmental approach to human evolution and sociality. In F. Jaroš, J. Klouda (Eds.), Adolf Portmann: A thinker of self-expressive life (pp. 89 – 118). New York: Springer.10.1007/978-3-030-67810-4_6Search in Google Scholar

Laland, K. N., Brown, G. R. (2018). 'The Social Construction of Human Nature', in Elizabeth Hannon, Tim Lewens (Eds.), Why We Disagree About Human Nature. Oxford: Oxford University Press.10.1093/oso/9780198823650.003.0008Search in Google Scholar

Laland, K. N., Galef, B. G. (Eds.) (2009). The question of animal culture. Cambridge, MA: Harvard University Press.10.2307/j.ctv322v4wfSearch in Google Scholar

Laland, K. N., Hoppitt W. (2003). Do animals have culture? Evolutionary Anthropology: Issues, News, and Reviews 12, 150 – 159.10.1002/evan.10111Search in Google Scholar

Leavens, D. A. (2004). Manual deixis in apes and humans. Interaction Studies: Social Behaviour and Communication in Biological and Artificial Systems, 5(3), 387 – 408.10.1075/is.5.3.05leaSearch in Google Scholar

Lewens, T. (2017). Human nature, human culture: the case of cultural evolution. Interface Focus 7: 20170018.10.1098/rsfs.2017.0018Search in Google Scholar

Lorenz, K. Z. (1981). The Foundations of Ethology. Vienna: Springer.10.1007/978-3-7091-3671-3Search in Google Scholar

Matsuzawa, T., Biro, D., Humle, T., Inoue-Nakamura, N., Tonooka, R., Yamakoshi, G. (2001). Emergence of culture in wild chimpanzees: Education by master-apprenticeship. In T. Matsuzawa (Ed.), Primate origins of human cognition and behavior (pp. 557 – 574). Tokyo: Springer.10.1007/978-4-431-09423-4_28Search in Google Scholar

McGrew, W. C., Tutin, C. E. G. (1978). Evidence for a social custom in wild chimpanzees? Man, 13, 234 – 251.10.2307/2800247Search in Google Scholar

Michelini, F. (2020). The philosopher's boredom and the lizard's sun: Martin Heidegger's interpretation of Jakob von Uexküll's Umwelt theory. In: Francesca Michelini, Kristian Köchy (Eds.): Jakob von Uexküll and Philosophy. London, New York: Routledge, pp. 122 – 140.10.4324/9780429279096-8Search in Google Scholar

Morand-Ferron, J., Cole, E. F., Rawles, J. E., Quinn, J. L. (2011). Who are the innovators? A field experiment with 2 passerine species. Behavioral Ecology 22, 1241 – 1248.10.1093/beheco/arr120Search in Google Scholar

Morrison, R. E., Eckardt, W., Colchero, F., Vecellio, V., Stoinski, T. S. (2021). Social groups buffer maternal loss in mountain gorillas. eLife, 10, e62939.10.7554/eLife.62939Search in Google Scholar

Olteanu, A. (2022). Learning as Becoming Conscious: A note on Jablonka and Ginsburg’s Notion of Learning. Biosemiotics 15, 457 – 467.10.1007/s12304-022-09510-xSearch in Google Scholar

Plessner, H. (1975). Die Stufen des Organischen und der Mensch. Berlin, New York: de Gruyter.10.1515/9783110845341Search in Google Scholar

Portmann, A. (1953). Das Kind vom Standpunkt der Anthropologie. Kölner Zeitschrift für Soziologie, 6, 441 – 446.Search in Google Scholar

Portmann, A. (1964). Das Tier als soziales Wesen. Freiburg i. Breisgau: Herder.Search in Google Scholar

Portmann, A. (1968). Die Biologie und das Phänomen des Geistigen. In A. Portmann, Biologie und Geist (pp. 9 – 49). Frankfurt a. M.: Suhrkamp.Search in Google Scholar

Portmann, A. (1990a). A zoologist looks at humankind. New York: Columbia University Press.Search in Google Scholar

Portmann, A. (1990b). The path leading to words: Levels of living communication. In Essays in philosophical zoology by Adolf Portmann: The living form and the seeing eye (pp. 97 – 124). Lewiston NY: The Edwin Mellen Press.Search in Google Scholar

Portmann, A. (1990c). Human beings in the perspective of the theory of evolution. In Essays in philosophical zoology by Adolf Portmann: The living form and the seeing eye (pp. 125 – 144). Lewiston NY: The Edwin Mellen Press.Search in Google Scholar

Reed, C. M. (2011). 'Till the Peoples All Are One'. Darwin's Unitarian Connections. London: Lindsey Press.Search in Google Scholar

Richerson, P. J., Boyd, R. (2005). Not By Genes Alone: How Culture Transformed Human Evolution. University of Chicago Press.10.7208/chicago/9780226712130.001.0001Search in Google Scholar

Rosenberg, K. (2021). The evolution of human infancy: Why it helps to be helpless. Annual Review of Anthropology, 50(1), 423 – 440.10.1146/annurev-anthro-111819-105454Search in Google Scholar

Stamm, R. A. (1999). L'intériorité, dimension fondamentale de la vie. Revue Européenne Des Sciences Sociales, 37(115), 55 – 73.Search in Google Scholar

Sterelny, K. (1996). Explanatory Pluralism in Evolutionary Biology. Biology and Philosophy 11, 193 – 214.10.1007/BF00128919Search in Google Scholar

Švorcová, J. (2023). Transgenerational Epigenetic Inheritance of Traumatic Experience in Mammals. Genes, 14, 120.10.3390/genes14010120Search in Google Scholar

Švorcová, J., Kleisner, K. (2018). Evolution by Meaning Attribution: Notes on Biosemiotic Interpretations of Extended Evolutionary Synthesis. Biosemiotics, 11, 231 – 244.10.1007/s12304-018-9328-9Search in Google Scholar

Švorcová, J., Markoš, A. (2019). Epigenetic processes and evolution of life. CRC Press.Search in Google Scholar

Tomasello, M. (1999). The Cultural Origins of Human Cognition. London: Harvard University Press.10.4159/9780674044371Search in Google Scholar

Tomasello, M., Savage-Rumbaugh, S., Kruger, A. (1993). Imitative learning of actions on objects by children, chimpanzees and enculturated chimpanzees. Child Development, 64, 1688 – 1705.10.1111/j.1467-8624.1993.tb04207.xSearch in Google Scholar

Whiten, A. et al. (1999). Cultures in chimpanzees. Nature, 399, 682 – 685.10.1038/21415Search in Google Scholar

Whiten, A. (2017). Culture extends the scope of evolutionary biology in the great apes. PNAS 114. No. 30, 7790 – 7797.10.1073/pnas.1620733114Search in Google Scholar

Wild, M. (2021). Portmann, Goethe and modern biology: Two and a half ways of looking at nature. In F. Jaroš, J. Klouda (Eds.), Adolf Portmann: A thinker of self-expressive life (pp. 145 – 158). New York: Springer.10.1007/978-3-030-67810-4_8Search in Google Scholar

Wilson, E. O. (1995). The Naturalist. Harmondsworth: Penguin.Search in Google Scholar

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