Parameters of Animal Self-Medication Behavior
By comparing self‑medication and caregiving among animals with evidence from the fossil record, we can reinterpret the deep‑time origins of self-medication behavior and recognize treating the injured and ill as a shared biological heritage rather than a uniquely human achievement.
Introduction
The Latin phrase Vis medicatrix naturae means “the healing power of nature.” The concept is attributed to Hippocrates, the ancient Greek physician regarded as the father of medicine. While he might have understood the restorative power of being in nature, one wonders whether he would be astounded by what we have learned from nature and what we still struggle to comprehend.
From the leaves that crown the tallest trees to the small seedlings tucked beneath the shelter of shrubs, the healing properties of plants are spread widely. Mother Nature, it seems, is a good physician. This power of healing can also be found across the animal kingdom, with animals (vertebrate and invertebrate) engaging in behaviors that resemble treatment and prevention against illness and disease. This behavior is called self‑medication. Research is shedding light on this act of healing, making us reconsider what we view as a uniquely human characteristic. The art and science of medicine is an area of social behavior where humans have formalized healing practices into a structured process. Yet the roots of our collective knowledge about healing stretch far deeper, forming a continuum that begins with animals. Whether driven by instinct or the will to survive, we can view self-medication behavior as a gradient stretching from simple acts to more formal, structured procedures for treating the ill and curing the sick. Yet the consensus is that a clear distinction needs to be made between accidental and instinctual behavior, which has set the stage for human-initiated formal medical procedures.
We know that healing from nature is older than humanity. Evidence of self-medication can be found among nonhuman communities in the past and present. But not all self-medication behavior is equal. Some behavior is natural, almost instinctive, while other behavior is socially learned, culturally transmitted, and context‑dependent. By comparing self‑medication and caregiving among animals with evidence from the fossil record, we can reinterpret the deep‑time origins of self-medication behavior and recognize treating the injured and ill as a shared biological heritage rather than a uniquely human achievement. Caution is, however, needed when interpreting evidence of such behavior in the wild and in the fossil record.
In this article, I explore evidence supporting the range of parameters for self-medication behavior to gain a more concrete understanding of the distinction between simple acts and more intentional behavior. As I dive into this discussion, it is important that self-medication behavior be viewed as a continuum rather than a single category.
Self-Medication Behavior: Getting to the Root of Origin
Humans and animals have used self-medication as a biological toolkit. Accounts of humans treating the sick and tending to the injured are widely documented in the archaeological record and include examples such as the ancient Egyptian Ebers Papyrus, a medical scroll, (c. 1550 BCE), the discovery of surgical instruments from ancient Rome (first century BCE to fourth century CE), and Chirurgia Practica by Roger of Salerno or the practice of surgery (c. 1180 CE). Early accounts of animal self-medication behavior were recorded by Aristotle in Historia animalium (384–322 BCE) and Pliny the Elder (c. 77–79 CE) in Naturalis historia. While humans relied on accumulated knowledge, cultural learning, and increasingly formalized techniques, animals have depended solely on nature. For them, the cure for many ailments is found in the “pharmacy of the wild.”[1]
In 2024,[2] a Sumatran orangutan from Indonesia made headlines globally after the publication of a study citing a 2022 incident. A research team from Germany and Indonesia, who were documenting the orangutans of Suaq Balimbing, noted a male orangutan, called Rakus, with a fresh wound on his face. The injury was most likely due to an altercation with another male. What surprised the team, and the world, was Rakus’s treatment of the wound. After chewing some leaves of a climbing shrub (Fibraurea tinctoria), he applied the chewed leafy paste to his face with his fingers, covering the wound entirely. When the team saw Rakus again a few days later, the wound was closed, and a month after, it had fully healed. Rakus’s behavior stunned and interested many people because he demonstrated a targeted, repeated, and seemingly intentional treatment of the wound. This discovery prompted the world to reconsider the possibility of a deeply rooted knowledge across species for using nature and its substances as part of the healing process. Rakus’s story might be the most well-known, but similar accounts exist for other animals.
This type of behavior is called zoopharmacognosy. It was first coined by Eloy Rodriguez and Richard Wrangham[3] to refer to the use of medicinal plants and their pharmacological properties by animals to treat injuries and discomfort. This concept has since expanded to include the use and consumption of minerals and other substances to treat injuries, diseases, and infections.[4][5] Zoopharmacognosy mainly focuses on the self-directed medication behavior of an individual toward itself.
Two types of self-medication behavior have been identified in animals, curative and preventative.[6] [7] The first is related to treating the body to maintain good health.[8] In this category, the animal’s behavior is indicative of whether it is ill. Slight changes in the animal’s diet are also noted. Specifically, if ill, non-nutritional food sources are consumed in small quantities often and for a limited period. Parts of specific plants are eaten or chewed in an attempt to cure the individual of a disease or its symptoms. Preventive action, on the other hand, is behavior that helps to prevent infection and can even be defensive. This can be coupled with a behavioral change to avoid the spread of the disease, such as avoiding contaminated food or water and even ill members of a group.[9] It can also include the ingestion of non-nutritional substances as a defense against future infection or parasites, like the monarch butterfly eating milkweed (Asclepias) so birds don’t feed on them.[8]
The challenge of identifying self-medication behavior in animals is that it requires a distinction between consuming plants or other substances having nutritional value and those eaten intentionally to relieve discomfort. This differentiation is required as some plants contain secondary metabolites such as alkaloids, tannins, and terpenoids, which are naturally used by the plant as defense against herbivores, pests, and environmental stress.[10] These secondary metabolites can also help fight off parasites. Meaning that some medical benefits from consuming a plant may be accidental rather than deliberate. The distinction between accidental and deliberate self-medication behavior, therefore, hinges on whether the animal is choosing specific plants or substances intentionally because there is a known link between the substance and feeling better.[8]
Both animals and humans look for solutions to ease pain and discomfort when sick. Instincts might have driven us to find solutions for healing in nature, but humans have formalized this process through medicine. Animals, meanwhile, somehow retain this knowledge and information about where to seek help and in what quantity. But for considering the behavior of using plants as part of a self-medication regime, the plants should have no food or nutritional value and be naturally toxic, bitter, or poisonous.[11]
This behavior leads to a further point of contention. If the behavior is truly self-medication, it needs to be taught or learned. This means even in animals, some form of knowledge transfer or exchange has occurred. [7][12] There are a few instances that can help us understand how knowledge can be passed between conspecific groups. First, knowledge can be exchanged and learned from females who transfer into new social groups. Behavior related to sourcing food, fighting predators, and treating an injury or infection can be shared (naturally or intentionally). Groups in the same region can also observe and learn behavior from each other.[13] The bond between mother and infant also allows for the transfer of knowledge to occur.[14] The last category includes the discovery of knowledge by using what is available until something works and retaining that knowledge for future application. This is how humans developed as pharmacists and doctors by learning what works through trial and error, observing animal behavior and other communities, and by sharing and exchanging knowledge with related and neighboring groups. What is clear in animal self-medication is that the behavior is targeted and not random.
Self-medication behavior can be self-directed (actions taken by an individual that benefit themselves) or other-directed, which are actions taken by an individual to benefit other individuals.[15] When medication behavior is directed toward others, it can either be individuals who are close kin, non-kin but part of the community, or strangers from other kin groups.[15] Other-directed self-medication reflects prosocial tendencies, as observed in a 2022 study where three cases of chimpanzees treating other members’ wounds through application of insects were reported.[16] A second study from 2025 also reported on one case where the use of insects in prosocial wound treatment by chimpanzee groups from Uganda was observed. [17] The latter study also reported that the transmission of insect use in wound care is learned through observation and social bonding. Both studies show that even in isolated regional contexts, prosocial care is inherent in primate species and could potentially be a behavior that existed before the split in our family tree.
The variation in behavior reported in the animal kingdom suggests that it might be driven beyond just the need to ease pain and discomfort. Reasons for this behavior include survival. The strongest survive, and the healthiest grow old. This also means the stronger and healthier individual is in a better position to attract a mate and produce strong and healthy offspring. This behavior, and perhaps the natural and learned instinct, are then driven by biological need. Individuals may select different care strategies based on symptoms, wound or infection severity, location, environmental context (including plant availability), or prior experience with particular treatment methods.[15] In the fight against parasites and pathogens, and to ensure survival, animals leverage the resources they have in abundance.
Types of Animal Self-Medication Behavior
The methods and strategies used by animals to treat maladies are as diverse as the types of biological invaders they need to fight. We can understand and interpret self-medication behavior through a six-point framework. This is not a final or exhaustive model, but it provides a backbone to interpret animal behavior, especially to understand whether the behavior is accidental or intentional, and if it is for health or survival purposes.
Symptoms and situations: Self-medication behavior is triggered by a specific problem, such as a parasite, infection, or injury. Depending on the symptoms or circumstances, the behavior can either be curative or preventive. The behavioral attribute, plant consumption, or substance use increases or only occurs when the animal is ill or hurt, or after it shows specific symptoms.
Expected medical behavior: Animals exhibit a range of behaviors to fight against pathogens and parasites. They are mainly grouped into pith chewing, leaf swallowing, plant eating, fur rubbing, and geophagy. This includes using plants and other substances that are non-nutritious. Importantly, the behavior and usage only take place when the animal is in discomfort. The behavior also differs from the accepted normal behavior and diet for the animal.
Localized specificity: The substance, which may be a plant or something else, is applied and used to target a specific problem or condition. Additionally, the substance or plant has a known pharmacological property, making it effective for treating specific injuries or infections, or as a preventative measure.
Non-nutritional: It is important for the behavior to have a functional outcome, specifically to improve an individual’s health and, as a result, their survival. The substances and plants are consumed purely for treatment and are not part of a regular diet or for hydration.
Dosage: In addition to the animal taking specific actions for treatment and applying substances and plants to targeted areas, the dosage of the application is also important. Knowledge of the exact amount and frequency of application indicates a potential therapeutic goal.
Socially instilled: The behavior is adopted by an animal after either observing other animals use the substance for treating injuries or infections, or is learned through social bonding.
To distinguish between intentional and accidental behavior, researchers have identified a list of practices that can be interpreted, within the above framework, as indicative of self-medication behavior. The behavior is classified into five categories.
Pith Eating
Bitter pith eating has been observed mostly in chimpanzees.[4][18] The inner stem of the bitter leaf plant (Vernonia amygdalina) is chewed, and the juices are swallowed. Leaves and the outer bark are stripped from the plant to get to the soft inner part. Chewing pith helps fight parasitic infestations (like nodular worms), stomach pain, and diarrhea. This targeted behavior is used to treat symptoms that especially appear to flare up during the rainy season.[18] Young chimpanzees learn this behavior by observing their mothers or other group members.[19]
Leaf Swallowing
Swallowing whole leaves without chewing them first is another therapeutic behavior that has been reported among chimpanzees, bonobos, gorillas, snow geese, and brown bears.[18] Similar to pith eating, this behavior is learned by younger generations, who observe and copy their older group members. Whole leaves are consumed by placing them in the mouth, folding them with the tongue, and then swallowing them without chewing.[20] Field researchers have observed expelled leaves containing parasites (tapeworm or nematodes) from individuals who swallowed them whole.[21] [22] It is thought that the structure and chemical compounds of specific leaves (like Aspilia) aid in trapping and removing worms as they pass through the gut.[4]
Plant Eating
Animals pick specific plants to chew on to ease discomfort. It is used to treat parasitic infections and other bacterial infections with the antimicrobial properties of plant substances. This self-medication behavior has also been reported in invertebrates, for example, caterpillars eat hemlock leaves when infected with parasites.[23] This behavior can also be used to support other health and well-being needs. An elephant from the Tsavo Park, Kenya, was recorded eating a borage tree to help induce labor[24] while lemurs chew on tamarind and fig leaves to increase milk production.[25]
Fur Rubbing
Within the broader self-medication category, fur rubbing can serve as both preventive and defensive behavior. To fend off parasites, fungal or bacterial infections, some animals rub their fur or part of their bodies with different substances. Some use the leaves or fruit of plants and apply them as a topical ointment, like a natural repellent remedy. Fur rubbing or self-anointing has been observed in capuchin monkeys (Cebus capucinus) who use shrubs (Piper marginatum) to expel insects and reduce infections, and in a group of spider monkeys (Ateles geoffroyi) who rub strong-smelling leaves on their bodies.[26] [27] [28] [29] [4] Wild Kodiak and brown bears use a different strategy. They chew asha root (Ligisticum porteri) and then spread their saliva mixed with the root into their fur. [4] Through this self-anointing behavior, they not only expel insects but also reduce skin irritation resulting from parasitic bites. White-nosed coatis (Nasua narica) found on Barro Colorado Island, Panama, take the menthol-like smelling resin from freshly scraped bark of Trattinnickia aspera (Burseraceae) and vigorously rub it into their own fur.[30] Animals are resourceful. Consuming citrus has huge nutritional benefits. But when applied to external surfaces of the body, it serves an entirely different purpose. Animals have been recorded applying the peel and fruits of citrus plants in a curative or preventative manner to reduce irritation. The chemical compounds found in citrus plants also repel insects (like ticks) and have been used for self-anointing and fur-rubbing by monkeys and birds.[31]
In other cases, substances other than plants have also been used. In Venezuela, capuchin monkeys (C. olivaceus) rub millipedes onto their fur to repel insects.[32] Ring-tailed coatis from Brazil have adapted to self-anointing using soap left outside. They rub bitten and chewed pieces of soap on their bodies and tails with their hands. The use of soap was the result of opportunistic exploration of human substances by coatis. And after establishing it as a useful repellent of parasites, the behavior was most likely transferred through social learning or imitation within the ring-tailed coati groups.[33] In a study from Costa Rica, a white-nosed coati was recorded rubbing the fecal boli of a male tayra on its tail.[34] Fur rubbing can thus be implemented as a survival strategy to fend off other predators by masking their own scent. Pandas, on the other hand, rub fresh horse manure on their bodies.[35] Observed more frequently in colder seasons, the chemical compounds in the manure block certain nerve signals in the skin, which help the pandas better cope with the cold. In warmer months, when drinking water is scarce, baboons from the Namib desert in Namibia regulate their body temperature by flicking plumes of sand from shady areas over their bodies (sandbathing). [36] Fur rubbing can take many forms. But what is clear is that it is an intentional behavior directed toward health and survival.
Geophagy
Unless you are an earthworm, eating soil is not a typical part of the mammalian diet. In animals, eating soil, or geophagy, reflects an underlying health concern. Elephants from the Udawalawe National Park, Sri Lanka, eat sodium-rich clay to detoxify their systems from previous substances consumed and to replenish minerals lost during dry seasons,[37] while elephants from Ugalla River Game Reserve, Western Tanzania, consume soil from termite mounds, which are rich in minerals.[38]
Japanese macaques, rhesus macaques,[39] mountain gorillas, chimpanzees,[40] and cattle also share a preference for sodium-rich soils.[15] The chemical and mineral composition of soil is thought to maintain gut pH, satisfy hunger for sodium, and detoxify the body from previously consumed toxic non-nutritional sources, and can also act as a cure for upset stomachs and diarrhea.[41] While geophagy peaks in the dry seasons to replenish lost minerals, it is also interesting that its use shifts in the wet season, where it is mainly used as digestive relief.[30]
Licking and Dabbing
Animals clean wounds by licking the injured areas. Pet owners are familiar with this behavior, often seen in their dogs. Chimpanzees, on the other hand, use their fingers to smear saliva on a wound.[15] Saliva is used as a self-soothing tactic and could be more instinctual than intentional medicinal behavior. But it does function to remove some debris from the wound. Another tactic to remove debris from wounds or stop bleeding is by dabbing leaves over the wound. This prevents further infection, most likely attributed to the antimicrobial and anti-inflammatory properties of the plants.[15]
Self-soothing behavior can be a reflection of an animal’s need for comfort during periods of pain. It can be instinctual. But if we consider this behavior in combination with wound cleaning using non-nutritional plants or other objects (twigs), it is more suggestive of behavior that leans toward self-medication. In addition to using vegetation as part of self-medication behavior, primates expand their first aid kit by using other substances (insects) in wound care strategies. Chimpanzees have been reported to treat wounds on themselves and others by dabbing and reapplying insects on injured areas as a topical treatment.[16]
Archaeological Evidence of Hominin Medical Behavior
We recognize self-medication behavior in animals due to similar behavioral patterns in human history. It’s not surprising because some of our own medical knowledge and use of plants stems from observing animals. However, recognizing a plant’s effect on yourself is very different from applying that knowledge to treat others, which requires more complex cognitive ability and establishing lasting social relationships and trust.
This distinction becomes important when interpreting evidence from the past. While we can trace humanity’s exploration of medical behavior in the archaeological record, one major issue is that plant material rarely survives. This makes it nearly impossible to determine with certainty whether the plant material that did survive was used medicinally. However, while we can’t rely on the survival of plant material, archaeologists and paleontologists can use other available sources to closely look at the skeletal material of our ancestors and observe their injuries. When excavating new archaeological sites, skeletal material preserves evidence of injuries and also the healing process. Healed fractures, amputations, and trepanation are examples of medical procedures that have been preserved in the archaeological record.
The earliest evidence of long-term survival after an injury or illness was found in Dmanisi, Georgia, dating 1.77 million years ago.[42] The individual (Homo erectus) lived and survived with periodontal disease: an infection of the gums and supporting tissues that can lead to inflammation of the jaw, bone loss, and, in severe cases, tooth loss.[43] This individual suffered the loss of chewing function, indicating that they probably had to adapt their diet and could have been reliant on the community for social support. This suggests that some forms of caregiving existed in early Homo groups.
Later in the Paleolithic, burials excavated at Shanidar Cave, Iraq, preserved not only behavior related to mortuary treatment but also further evidence of social care and the treatment of the injured. Skeletal material of a Neanderthal male dating to approximately 45,000–70,000 years ago, indicates that before death, he survived with an unrepaired fracture of the right arm and lived until around 40 years old.[44] [45] Other skeletal remains from Shanidar Cave[46] reveal that these individuals not only received treatment for their injuries but also survived long enough for the injuries to fully heal. Their later deaths were unrelated to their injuries. The treatment of these individuals most likely included the use of plant substances. Interestingly, pollen clusters of plants were excavated near the graves of these individuals, possibly indicating evidence of intentional placement over or near the deceased.[47] [48]
Molecular compounds from common yarrow (Achillea millefolium) and chamomile (Matricaria chamomilla) were found on Neanderthal remains buried at El Sidrón, Spain. These compounds were extracted from dental buildup (dental calculus) from an individual who lived approximately 49,000 years ago.[49] While an interesting and unexpected find, it is difficult to prove that past communities used plant substances as part of their treatment regimen, as there is no way to confirm the intentions or purpose behind the inclusion of these plants. The El Sidrón skeletal material, however, revealed extensive information about the individual’s health. Analysis of the gut microbiome and a tooth abscess, paired with molecular samples, indicates consumption of non-nutritional plant substances (yarrow and chamomile), showing some knowledge and suggesting a form of proto-medicine (pre-scientific and traditional healing approaches) within the social group.[50]
Medical behavior is not limited to Neanderthals. Examples of medical behavior have also been uncovered for Homo sapiens. Skeletal material from Liang Tebo Cave, Borneo, provided information about a young individual who survived six to nine years after a lower limb amputation dating to 31,000 years ago.[51] Later in the Upper Paleolithic, from the Villabruna burial, located in the Riparo Villabruna in Sovramonte, Belluno, northern Italy, evidence of invasive dental treatment survived. The dental remains of a young male preserved signs of dental treatment expressed through scrape marks found on the inside of his teeth.[52] The earliest trepanation of a 50-year-old male is from Vasilyvkil, Ukraine, dating to the Mesolithic 8020–7620 years ago.[53] Together, these examples demonstrate that the communities of the Later Paleolithic relied on their kin for food, support, and treatment when they were injured or ill.
The examples above indicate that the roots of healing and treatment extend far into human history. Examples from both animal and fossil records show that this behavior stemmed from a personal need to ease pain and an evolutionary instinct to survive. But when did we make the jump from genetically driven behavior to a more formal, socially organized system for treating the injured and sick?[11] A possible answer could be that for medicine to emerge, a species needs to possess both cognitive abilities and social structures that support intentional treatment. This is similar to moving from mortuary behavior to the intentional burial of kin, which requires communities to possess two key qualities: complex cognitive abilities and socially organized structures for intentional treatment.
In terms of cognition for self-medication, this boils down to recognizing symptoms and their potential causes and finding and applying the right treatment in a targeted manner. This knowledge, as far as researchers can determine, remains within the memory of social groups and is transmitted through observations and kin teaching. On the other hand, treating the sick and injured also requires social structures for establishing stable social groups and caregiving norms. Groups, therefore, need to have the capacity socially, economically, and mentally to take care of the injured and sick.[54] This includes establishing long-term bonds in the community.
Finally, a knowledge system with shared, traditional, or Indigenous know-how about plant application and use is transmitted across generations. Combined, these points show that medicine emerges not from sudden innovation but from the interaction of social care and environmental knowledge. The move from self-medication behavior to formal medicine can then be viewed as a spectrum, rather than a linear progression. For this to occur, three things need to happen in a group, according to a 2004 unpublished paper by Steven Cartwright[11]:
The use of plants and other substances for the treatment of injuries and infections.
Ritualized behavior related to healing (also referred to as shamanism).
Knowledge and development of processes and tools used in physical procedures.
Groups with socially organized medical systems have taken advantage of these three areas. Animals exhibit self-medication behavior, while Neanderthals and Homo sapiens used a combination of ritualized healing and proto-medicine. Meanwhile, modern humans formalized these healing structures into an organized social system.

Returning to the Roots of Medicine
The parameters that comprise the gradients of self-medication behavior provide a baseline through which we can interpret and understand animal behavior, and maybe our past selves. Animals heal themselves and one another with intention, memory, and skill, and early hominins continued and expanded these behaviors long before formal medical systems appeared. The parameters provide a lens through which we can view past behavior. The archaeological record shows that Neanderthals and early Homo sapiens recognized illness, and in certain circumstances, used medicinal plants, cared for the injured, and even performed complex procedures. This behavior isn’t static. It’s a dynamic conceptual model that needs to be updated as we learn more about ourselves and the species around us.
In animals, and possibly in the past, self-medication should be viewed as more than a binary system. By mapping the parameters that separate the different phases, we avoid the trap of overinterpreting simple accidental behavior as intentional. Instead, when we view it as a gradient of medical possibility, it opens a landscape where animals and early hominins experimented with healing in diverse ways. These practices reveal that medicine is not a recent cultural invention, but is an ancient evolutionary strategy rooted in social cooperation, environmental knowledge, and a biological need to survive.
The closer we look, the more we realize that the pharmacists of the wild are showing us that we have much to learn about the natural world. In fact, there might be some forgotten natural treatment locked up in Indigenous and traditional knowledge, which modern science and technology may yet rediscover. In this sense, one could argue that self-medication behavior and medicine are not human inventions but our shared inheritance.
- ↑ R.J. Safran (2025). “Pharmacy of the Wild: Lessons from Animal Doctors.” Current Biology. Vol. 35, Issue 11, pp. R399–R400.
- ↑ BBC News (2024). “Wounded Wild Orangutan Seen Using Plant as Medicine.”
- ↑ E. Rodriguez and R. Wrangham (1993). “Zoopharmacognosy: The Use of Medicinal Plants by Animals.” In Phytochemical Potential of Tropical Plants. K.R. Downum; J.T. Romeo; and H.A. Stafford (eds.). New York: Springer. Pp. 89–105.
- ↑ 4.0 4.1 4.2 4.3 4.4 M.A. Huffman (1997). “Current Evidence for Self-Medication in Primates: A Multidisciplinary Perspective.” American Journal of Biological Anthropology, Yearbook of Physical Anthropology. Vol. 104, Issue S25, pp. 171–200.
- ↑ J.C. De Roode and M.A. Huffman (2024). “Animal Medication.” Current Biology. Vol. 34, Issue 17, pp. R808–R812.
- ↑ R. Raman and S. Kandula (2008). “Zoopharmacognosy: Self-Medication in Wild Animals.” Resonance. Vol. 13, Issue 3, pp. 245–253.
- ↑ 7.0 7.1 J.C. De Roode; T. Lefèvre; and M.D. Hunter (2013). “Self-Medication in Animals.” Science. Vol. 340, Issue 6129, pp. 150–151.
- ↑ 8.0 8.1 8.2 M.A. Huffman and S.K. Vitazkova (2007). “Primates, Plants, and Parasites: The Evolution of Animal Self-Medication and Ethnomedicine.” In Ethnopharmacology. EOLSS Publications.
- ↑ P.C. Lopes; S.S. French; D.C. Woodhams; and S.A. Binning (2022). “Infection Avoidance Behaviors Across Vertebrate Taxa.” Animal Behavior and Parasitism. V.O. Ezenwa; S.M. Altizer; and R.J. Hall (eds.). Oxford: Oxford University Press.
- ↑ E.S. Teoh (2015). “Secondary Metabolites of Plants.” Medicinal Orchids of Asia. Vol. 5, pp. 59–73.
- ↑ 11.0 11.1 11.2 S. Cartwright (2004). “The Origins of Medicine; Assessment and Implications of the Eurasian Evidence from the Upper Palaeolithic to the Bronze Age.” Unpublished paper.
- ↑ J. Shurkin (2014). “News Feature: Animals That Self-Medicate.” Proceedings of the National Academy of Sciences. Vol. 111, Issue 49, pp. 17,339–17,341.
- ↑ J. Duboscq; V. Romano; A. MacIntosh; and C. Sueur (2016). “Social Information Transmission in Animals: Lessons from Studies of Diffusion.” Frontiers in Psychology. Vol. 7, Issue 1147.
- ↑ J.J. Villalba and F.D. Provenza (2007). “Self-Medication and Homeostatic Behaviour in Herbivores: Learning About the Benefits of Nature’s Pharmacy.” Animal. Vol. 1, Issue 9, pp. 1,360–1,370.
- ↑ 15.0 15.1 15.2 15.3 15.4 15.5 E. Freymann; C. Hobaiter; M.A. Huffman; H. Klein; et al. (2025). “Self-Directed and Prosocial Wound Care, Snare Removal, and Hygiene Behaviors Amongst the Budongo Chimpanzees.” Frontiers in Ecology and Evolution. Vol. 13, Article no. 1540922.
- ↑ 16.0 16.1 A. Mascaro; L.M. Southern; T. Deschner; and S. Pika (2022) “Application of Insects to Wounds of Self and Others by Chimpanzees in the Wild.” Current Biology. Vol 32, Issue 3, pp. R112-R113.
- ↑ K. Kolff; D. Acosta Flórez; A. Mascaro; and S. Pika (2025). “Insect Applications to Open Wounds by Chimpanzees in the Wild: First Insights From East African Chimpanzees.” Scientific Reports. Vol. 15, 31242.
- ↑ 18.0 18.1 18.2 M.A. Huffman (2003). “Animal Self-Medication and Ethno-Medicine: Exploration and Exploitation of the Medicinal Properties of Plants.” Proceedings of the Nutrition Society. Vol. 62, Issue 2, pp. 371–381.
- ↑ J. Bray; M.E. Thompson; M.N. Muller; R.W. Wrangham; and Z.P. Machanda (2017). “The Development of Feeding Behavior in Wild Chimpanzees (Pan Troglodytes Schweinfurthii).” American Biological of Physical Anthropology. Vol. 165, Issue 1, pp. 34–46.
- ↑ M.A. Huffman; J.E. Page; M.V.K. Sukhdeo; S. Gotoh, et al. (1996). “Leaf-Swallowing by Chimpanzees: A Behavioral Adaptation for the Control of Strongyle Nematode Infections.” International Journal of Primatology. Vol. 17, Issue 4, pp. 475–503.
- ↑ M.A. Huffman and S. Hirata (2004). “An Experimental Study of Leaf Swallowing in Captive Chimpanzees: Insights Into the Origin of a Self-Medicative Behavior and the Role of Social Learning.” Primates. Vol. 45, Issue, 2, pp. 113–118.
- ↑ A. Fowler; Y. Koutsioni; and V. Sommer (2007). “Leaf-Swallowing in Nigerian Chimpanzees: Evidence for Assumed Self-Medication.” Primates. Vol. 48, Issue 1, pp. 73–76.
- ↑ C.R. Engel (2002). “Acknowledging the Potential Role of Animal Self Medication.” Proceedings of the UK Organic Research 2002 Conference. Organic Center Wales, Institute of Rural Studies, University of Wales Aberystwyth. Pp. 355–358.
- ↑ K.P.N. Shuker (2001). “The Hidden Powers of Animals: Uncovering the Secrets of Nature.” London: Marshall.
- ↑ M.L. Sauther (1994). “Wild Plant Use by Pregnant and Lactating Ringtailed Lemurs, With Implications for Early Hominid Foraging.” In Eating on the Wild Side: The Pharmacologic, Ecologic and Social Implications of Using Non-cultigens. N.L. Etkin (ed.). Tucson: University of Arizona Press (1994). Pp. 240–256.
- ↑ C.J. Campbell (2000). “Fur Rubbing Behavior in Free-Ranging Black-Handed Spider Monkeys (Ateles geoffroyi) in Panama.” American Journal of Primatology. Vol. 51, Issue, 3 pp. 205–208.
- ↑ M. Baker (1996). “Fur Rubbing: Use of Medicinal Plants by Capuchin Monkeys (Cebus capucinus).” American Journal of Primatology. Vol. 38, Issue 3, pp. 263–270.
- ↑ M. Baker (1996). “Fur Rubbing by Capuchin Monkeys.”
- ↑ M. Bowler; E.J.E. Messer; N. Claidière; and A. Whiten (2015). “Mutual Medication in Capuchin Monkeys—Social Anointing Improves Coverage of Topically Applied Anti-Parasite Medicines.” Scientific Reports. Vol. 5, Article no. 15030.
- ↑ 30.0 30.1 R.M. Holdø; J.P. Dudley; and L.R. McDowell (2002). “Geophagy in the African Elephant in Relation to Availability of Dietary Sodium.” Journal of Mammalogy. Vol. 83, Issue 3, pp. 652–664.
- ↑ P.J. Weldon; J.F. Carroll; M. Kramer; R.H. Bedoukian; et al. (2011). “Anointing Chemicals and Hematophagous Arthropods: Responses by Ticks and Mosquitoes to Citrus (Rutaceae) Peel Exudates and Monoterpene Components.” Journal of Chemical Ecology. Vol. 37, Issue 4, pp 348–359.
- ↑ P.J. Weldon; J.R. Aldrich; J.A. Klun; J.E. Oliver; et al. (2003). “Benzoquinones From Millipedes Deter Mosquitoes and Elicit Self-Anointing in Capuchin Monkeys (Cebus spp.).” Naturwissenschaften. Vol. 90, Issue 7, pp. 301–304.
- ↑ A. Gasco; A.M. Pérez-Acosta; and P. Ferreira Monticelli (2016). “Ring-Tailed Coatis Anointing With Soap: A New Variation of Self-Medication Culture?” International Journal of Comparative Psychology. Vol. 29, Issue 1, pp. 1–11.
- ↑ F. Fleming and P.J. Weldon (2021). “Report of White-Nosed Coati (Nasua narica) Rubbing Itself With Feces of Tayra (Eira barbara) in Costa Rica.” Therya Notes 2. Vol. 2, Issue 3, pp. 176–178.
- ↑ W. Zhou; S. Yang; B. Li; Y. Nie; et al. (2020). “Why Wild Giant Pandas Frequently Roll in Horse Manure.” Proceedings of the National Academy of Sciences. Vol. 117, Issue 51 pp. 32,493–32,498.
- ↑ C. Brain and D. Mitchell (1999). “Body Temperature Changes in Free-ranging Baboons (Papio hamadryas ursinus) in the Namib Desert, Namibia.” International Journal of Primatology. Vol. 20, pp 585–598.
- ↑ R. Chandrajith; E. Kudavidanage; H.J. Tobschall; and C.B. Dissanayake (2009). “Geochemical and Mineralogical Characteristics of Elephant Geophagic Soils in Udawalawe National Park, Sri Lanka.” Environmental Geochemistry and Health. Vol. 31, Issue 3, pp. 391–400.
- ↑ E. Kalumanga; D.G. Mpanduji; and S.A.O. Cousins (2017). “Geophagic Termite Mounds as One of the Resources for African Elephants in Ugalla Game Reserve, Western Tanzania.” African Journal of Ecology. Vol. 55, Issue 1, pp. 91–100.
- ↑ N. Klein; F. Fröhlich; and S. Krief (2008). “Geophagy: Soil Consumption Enhances the Bioactivities of Plants Eaten by Chimpanzees.” Naturwissenschaften. Vol. 95, Issue 4, pp. 325–331.
- ↑ R. Raman and S. Kandula (2008). “Zoopharmacognosy: Self-Medication in Wild Animals.” Resonance. Vol. 13, Issue 3, pp. 245–253.
- ↑ R. Krishnamani and W.C. Mahaney (2000). “Geophagy Among Primates: Adaptive Significance and Ecological Consequences.” Animal Behaviour. Vol. 59, Issue 5, pp. 899–915.
- ↑ A. Margvelashvili; M. Tappen; G.P. Rightmire; N. Tsikaridze; and D. Lordkipanidze (2022). “An Ancient Cranium From Dmanisi: Evidence for Interpersonal Violence, Disease, and Possible Predation by Carnivores on Early Pleistocene Homo.” Journal of Human Evolution. Vol. 166, Article no. 103180.
- ↑ D. Lordkipanidze; A. Vekua; R. Ferring; G.P. Rightmire; et al. (2005). “The Earliest Toothless Hominin Skull.” Nature. Vol. 434, pp. 717–718.
- ↑ J.J. Hublin (2009). “The Prehistory of Compassion.” Proceedings of the National Academy of Sciences. Vol. 106, Issue 16, pp. 6,429–6,430.
- ↑ P. Spikins; A. Needham; L. Tilley; and G. Hitchens. (2018). “Calculated or Caring? Neanderthal Healthcare in Social Context.” World Archaeology. Vol. 50, Issue 3, pp. 384–403.
- ↑ E. Trinkaus and M.R. Zimmerman (1982). “Trauma among the Shanidar Neandertals.” American Journal of Biological Anthropology. Vol. 57, Issue 1, pp. 61–76.
- ↑ Ralph S. Solecki (1975). “Shanidar IV, a Neanderthal Flower Burial in Northern Iraq.” Science. Vol. 190, Issue 4217, pp. 880–881.
- ↑ A. Leroi-Gourhan (1975). “The Flowers Found with Shanidar IV, a Neanderthal Burial in Iraq.” Science. Vol. 190, pp. 562–564.
- ↑ K. Hardy; S. Buckley; M.J. Collins; A. Estalrrich; et al. (2012). “Neanderthal Medics? Evidence for Food, Cooking, and Medicinal Plants Entrapped in Dental Calculus.” Naturwissenschaften. Vol. 99, Issue 8, pp. 617–626.
- ↑ L.S. Weyrich; S. Duchene; J. Soubrier; L. Arriola; et al. (2017). “Neanderthal Behaviour, Diet, and Disease Inferred from Ancient DNA in Dental Calculus.” Nature. Vol. 544, pp. 357–361.
- ↑ T.R. Maloney; I.E. Dilkes-Hall; M. Vlok; A.A. Oktaviana; et al. (2022). “Surgical Amputation of a Limb 31,000 Years Ago in Borneo.” Nature. Vol. 609, pp. 547–551.
- ↑ G. Oxilia; M. Peresani; M. Romandini; C. Matteucci; et al. (2015). “Earliest Evidence of Dental Caries Manipulation in the Late Upper Palaeolithic.” Scientific Reports. Vol. 5, Article no. 12150.
- ↑ M.C. Lillie (1998). “Cranial Surgery Dates Back to Mesolithic.” Nature. Vol. 391, pp. 854–854.
- ↑ R. Dunbar (1996). “Grooming, Gossip, and the Evolution of Language.” U.S. Harvard University Press.

