Ethnobotanical inventory of medicinal plants used by Cashinahua (Huni Kuin) herbalists in Purus Province, Peruvian Amazon

We documented 467 plant taxa used for medicinal purposes by the Cashinahua community. In this paper, we present the general findings related to these 467 taxa with a main focus on 79 vascular plant species for which there is no documented medicinal use.

Out of the 467 taxa, most botanical samples were identified to species level, 105 to genus level, 6 to family level, while 12 taxa remained taxonomically unidentified. All the documented species are named in the Hantxa Kuin language. Some species were collected several times to complete fertile samples or were introduced by informants under a different vernacular name. We can observe, from a botanical perspective, an over-differentiation [45] of 72 plant species which bear two to four different local names. In six cases, under-differentiation was observed in which the identical vernacular name corresponded to different botanical species.

The 467 documented taxa belonged to 99 botanical families, the most predominant of which were Acanthaceae with 35 species, followed by Piperaceae (28 spp.), Rubiaceae (27 spp.), Araceae and Bignoniaceae (20 spp. each). The species most collected and used by informants were Pseuderanthemum lanceolatum (Ruiz & Pav.) Wassh. with 38 UR, Leonia glycycarpa Ruiz & Pav. and Piper reticulatum L. with 30 UR each, Piper aduncum L., Uncaria tomentosa D.C., Tradescantia zanonia (L.) Sw., Abuta grandifolia (Mart.) Sandwith, Matisia cordata Bonpl., Pseuderanthemum congestum (S.Moore) Wassh., Drymonia tenuis (Benth.) J.L.Clark, Mascagnia eggersiana (Nied.) W.R.Anderson and Piper leucophaeum Trel.

The most reported therapeutic activities were envenomation, particularly venomous snake and insect bites; infections and infestations, including herpes and conjunctivitis; the treatment of pregnancy and birth disorders, with the most prevalent uses being prenatal care, facilitating childbirth and accelerating labour; and digestive system disorders, for which the most frequently cited symptoms were diarrhoea, vomiting and constipation, and dental care. Almost all the taxa used in the treatment of culture-bound syndromes—unspecified socio-cultural use, family planning and religious use—also had normal therapeutic or prophylactic uses.

The most common forms of application were via external use, at 85.9%. Warm baths were the most prevalent (762 UR), which together with cold baths (89 UR) accounted for a total of 32.7% of the applications for external use. Direct application on the affected body part amounted to 34.4%, rubbing 15.1% and washing of the affected part 8.4%. The instillation of the juice of fresh leaves directly into the eye represented 5.6%. Oral administration accounted for 14.1% of the applications or 444 UR, which included the use of potions prepared by macerating pounded fresh plant material in water (64.6%, 287 UR), or decoctions (27%, 120 UR). Lastly, 1.1% of the recorded plant material, mostly leafy branches of Couepia obovata Ducke together with the leaves of the fern Pityrogramma calomelanos (L.) Link, burnt as a fumigant, was used to control infectious disease epidemics and to repel insects. The majority of our reported species (99%) were considered as wild, and they were usually collected in the nearby forest, along river banks and less often in anthropic environments (chacras) over quite a large gathering distance (5 km) from human settlements. In order to be ready for any emergency, each of the plant specialists maintained a small forest garden near the village. There, they transplanted some rare medicinal plants that are uncommon in the vicinity. Male plant specialists kept tree seedlings growing from natural reproduction around their paths well pruned so that they always had access to fresh leaves when needed. For the same reason, most samples collected from the trees were juveniles. Leaves were the most common plant part used (97%), explaining why the Cashinahua call their traditional medicine “dau pei”, leaf medicine.

Predominant families and species

From a total of 467 medicinal taxa collected, there are 99 different botanical families represented in our research, including Acanthaceae, Piperaceae, Rubiaceae, Araceae and Bignoniaceae which together represent more than one quarter of the total number of plants cited (Fig. 2). The situation is similar in other parts of the Ucayali region in which ethnobotanical studies have been carried out among Panoan groups [15, 20].

Fig. 2figure 2

Predominant botanical families. Only the 20 most common families (represented by 6 or more species) are shown

Representatives of the family Acanthaceae—three species of Pseuderanthemum: P. lanceolatum—mikin medan putani bata were used, mainly for pregnancy and birth disorders, infections manifesting as various forms of herpes and the treatment of upper respiratory tract inflammation. These species, as well as another two species of Pseuderanthemum, were used as first aid for venomous snake bites. Another representative of this genus, xuke bibex bata—P. congestum (S.Moore) Wassh., was used to treat herpes including shingles and cold sores (12 URs), venomous snake bites (6 UR) and eye problems such as cataracts. Dunu himi—Pseuderanthemum sp.—was used in 10 cases to treat venomous Bothrops atrox bites. Graham [15] also mentions Aphelandra lasiandra (Mildbr.) McDade & E.A.Tripp..—yawan kuxi dau—as one of the most widely used species in the Acanthaceae family, and this partially coincides with the use of this plant in our research. This plant’s spiny leaves are used to gently whip children’s legs to accelerate their walking. We documented the use of chewed leaves squeezed into the wound of venomous Scolopendra gigantea (Amazonian giant centipede) bites, in baths to strengthen a child’s body, in the treatment of headaches, and to treat strong convulsions identified by vegetalistas, or shaman, as the attack of a yawa (peccary) spirit or “peccary epilepsy”. According to Schultes and Raffauf [46], a toothache rinse can be made from an infusion of this plant. This plant’s root has been found to contain the alkaloid aphelandrine [47].

The Piperaceae family comprises 5 genera, of which Piper (about 2,000 species) and Peperomia (about 1,600 species) are the most important. The plants grow as herbs, vines, shrubs and trees and are widely distributed and used throughout the tropics and subtropics. Primary forests and forest edges, where we encountered the greatest number of useful species, are home to the greatest variety and abundance of Piper plants. Out of a total of 24 Piper taxa documented in our research, 14 were identified to species and 10 to genus. The most frequently used were Piper aduncum (31 UR), P. reticulatum (30 UR), P. leucophaeum (22 UR), and P. marginatum and P. peltatum L. (11 UR each). The Cashinahua people reported using P. aduncum leaves to treat digestive problems, pain, headaches, inflammation and fainting, as well as to prevent caries. P. reticulatum was primarily used for prenatal or childbirth care and to reduce high fever. Socio-culturally, it was used for behavioural regulation, to lower libido and for hunting. P. marginatum was mostly used to alleviate headaches, to cure inflamed teeth and to treat scorpion stings. Out of the 27 Piper species reported by Graham [15] there are two congruent species: yaix maxaka (P. nudilimbum) and txexen pei (P. peltatum). The first one is used to treat skin blemishes and mycotic infections, which coincides with our research. We documented the use of these two species in treating scabies, skin conditions and skin protuberances in the vagina or anus, genitourinary infections, and as a general restorative. In addition to the common use of P. peltatum in treating intestinal parasitosis, other uses such as healing stingray stings, cephalea (headache) and fainting were documented. In an indigenous Huni Kuĩ (Kaxinawa) community in the state of Acre, Brazil, adjacent to the current study area, Ehringhaus [13] gathered 48 species of Piper, of which 96% were used as analgesics, oral hygiene products and dermatological treatments. The antibacterial and antifungal properties of ethanol extracts of Piper spp. leaves and twigs have been examined. Around 44–60% of the extracts inhibited dermatophytic fungal growth, while 69–91% of the extracts inhibited bacterial growth [13]. Terpenoids, alkaloids and phenolic compounds were found in spot tests for secondary plant metabolites. Another genus of Piperaceae was collected, namely Peperomia, for which we documented 3 species: Peperomia blephariphylla Trel. & Yunck., Peperomia swartziana Miq. and Peperomia pilosa Ruiz & Pav.

Among the family Rubiaceae (27 spp., 180 UR), the species with the most reported uses was Uncaria tomentosa L. with 29 UR. The most frequently documented uses included the treatment of gastritis, rheumatism, internal tumours and hernias. As to the data on the use of plants of the Rubiaceae family in traditional Cashinahua medicine, we agree with Graham [15] regarding the species kawa (Psychotria viridis Ruiz & Pav.), matsi kawa (P. carthagenensis Jacq.), ixkin tepekan bata (Randia armata (Sw.) DC.) and xuniwan (Geophila macropoda (Ruiz & Pav.) DC.). The species are identical, but apart from the use of P. viridis we report different uses. R. armata (25 UR) was used for the treatment of snake bites, respiratory infections and skin diseases. According to Zamora-Martínez and Pola [48], a snake bite is treated in Mexico by taking a mixture of aguardiente (alcoholic liquor) and the fruit and twigs of this plant. Our respondents reported 10 uses of G. macropoda to cure inflamed molars and gingival abscesses with mouthwashes mostly in the form of a decoction of leaves. Within the Rubiaceae family, Psychotria was the most commonly occurring genus in our study with a wide range of uses. We documented the use of 7 taxa: P. alba Ruiz & Pav., P. carthagenensis, P. ruizii Standl., P. viridis, and 3 specimens identified to the genus only. The use of P. viridis in the preparation of nixi pae (ayahuasca) was recorded by Graham [15]. The leaves of Psychotria species are widely recognized as admixtures of the hallucinogenic beverage ayahuasca [7]. The leaves of P. carthagenensis and P. viridis have been found to contain indole alkaloids and sterols [49, 50].

The 20 collected specimens of Araceae which made up part of this study represent at least seven genera and 20 species. Among them, the most prevalent genus was Philodendron with 9 species, the most frequent being P. ernestii Endl. and P. fibrillosum Poepp. No literature reports of ethnomedicinal use, biological activity testing or phytochemical analysis were found for this species.

The most predominant species of Bignoniaceae (169 UR) were Dolichandra unguis-cati (L.) L.G.Lohmann, Cuspidaria floribunda (DC.) A.H.Gentry, Tanaecium dichotomum (Jacq.) Kaehler & L.G.Lohmann, Jacaranda glabra (DC.) Bureau & K.Schum. and Parmentiera cereifera Seem. For this species, there are no pharmacological reports in the scientific literature, apart from that of Gachet et al. [51], who demonstrate the promising activity of J. glabra against the Plasmodium falciparum K1 strain.

The species most collected and used from other families were Leonia glycycarpa Ruiz & Pav. (Violaceae), Tradescantia zanonia (L.) Sw. (Commelinaceae), Abuta grandifolia (Mart.) Sandwith (Menispermaceae), Matisia cordata Bonpl. (Malvaceae), Drymonia tenuis (Benth.) J.L.Clark (Gesneriaceae) and Mascagnia eggersiana (Nied.) W.R.Anderson (Malpighiaceae).

We specifically aim to highlight that the selected species in our study, however, do not fully represent the Cashinahua knowledge of medicinal plants.

Mode of preparation

The predominance of decoctions (1,247 UR), mostly made by boiling plant material in water for topical application (baths, washes and poultices) and for oral ingestion, agrees with that reported in other neighbouring regions [20]. We observed that elderly people generally prefer to prepare their remedies as decoctions because they believe that the more time the plant material is in contact with water, the more effective it is. Our respondents reported several preparation methods including soaking the fresh plant material (usually leaves) in a large volume of cold water (771 UR), soaking ground/pounded/grated leaves in a small volume of cold water (303 UR), and squeezing fresh leaf juice directly on the affected part (251 UR). In the vast majority of cases, water is the vehicle for almost all oral and topic preparations. Without exception, fresh plant material is always used in the preparation of remedies. Two-hundred and fifty-six UR (8.1% of remedies) were prepared in the form of patarashca, the typical Amazonian method of preparing small fish wrapped in the leaves of mani pui (Calathea lutea (Aubl.) Schult.), lightly roasted near the fire. The same form of preparation is applied by the Cashinahua to slightly crushed fresh leaves, to which the seeds of maxe “achiote” (Bixa orellana L.) are often added, mostly for the treatment of different skin conditions. The wax covering the under surface of C. lutea leaves, known in the Amazon as cauassú, may serve as a biological defence against infections and herbivory [52]. We presume that when heated this wax may aid in the therapeutic effects of a remedy prepared in form of patarashca.

Mode of application

It is interesting to note that only about 14% of the remedies were taken orally, with the vast majority (85%) being applied topically, most frequently as a bath. The remaining 1% served as protection against epidemics in the form of a fumigant (Table 1). The significant preponderance of externally applied remedies versus internal administration may seem surprising, but it is not uncommon among Amazonian and Andean ethnic groups [53, 54]. Alexiades [40], in his dissertation on the traditional medicine of the indigenous Amazonian Ese Eja people, mentions that 70% of remedies consisted of external treatments which involved direct contact of the plant tissue with the affected body part. The skin offers an accessible and convenient site for the administration of medications. To this end, the field of transdermal drug delivery, aimed at developing safe and efficacious means of delivering medications through the skin, has garnered much time and investment with the continuous advancement of new and innovative approaches [55]. It should be considered that the conventional use of medications in the form of oral administration must overcome the first pass effect, where the active substance enters the digestive tract and undergoes metabolic changes in the liver, which greatly slows down the onset of action and also alters its effects. Kováčik et al. [56] claim that the clinical advantages of transdermal drug delivery over traditional administration methods are numerous. In addition, a transdermal drug delivery system has been accepted as a potential non-invasive route of drug administration [57].

Table 1 Modes of application of 467 taxa based on 3,154 Use Reports)

The predominant external application methods were warm and cold baths, which together accounted for a total of 32.7% of external uses. Baths were usually prepared in the form of a decoction or pounded leaves macerated in water. The direct application of ground or chewed leaves squeezed on the affected part (22.7%) was used frequently in the first stage of treatment of venomous bites, herpes and cold sores. Crushed plant material heated in the form of patarashca was used for skin and subcutaneous cellular tissue disorders, leishmaniosis, infections, inflammations and injuries. Rubbing with fresh or heated leaves ground in a small amount of water (16.2%) was most often applied to areas of inflammation, swelling and pain, but also as a second stage in the treatment of snake bites, as well as in pregnancy care and childbirth. Washing of the affected part (7.4%) was reported to be useful to relieve rheumatic pain, headache, abscesses, haemorrhoids, haemorrhages and irregular menstruation, as well as to cure flu, diarrhoea, inflammations and skin infections. Poultices and plasters (2.6%) of crushed and mostly heated leaves were applied in cases of lumbago, paralysis of the face, nervous tics, bruises, closed wounds and weaning—prevention of mastitis. A special form of application used by different Amazonian ethnic groups [15, 58] is an ocular administration—the instillation of crushed leaf juice directly into the eye (5.6%) which is used to treat conjunctivitis, sties, affections of sight and headaches. However, its most frequent use was during an episode of fainting, dizziness and strong convulsions (a yuxin attack) that healers compared with epilepsy, in addition to socio-cultural and magical uses such as sorcery and “panema”, that is, misfortune in hunting. Another common use was its application before going hunting in order to be able to see animals better in the shadows of the forest.

Oral delivery has the advantages of allowing for pre-determined doses, portability and patient self-administration. For these reasons, the oral route in Western culture remains the most convenient means of delivering medications [59, 60]. Ingestion was applied for a wide range of ailments, mainly for the treatment of digestive, genitourinary and respiratory problems, as well as socio-cultural uses such as behavioural regulation and the nixi pae (ayahuasca) ceremony.

Drug activities

Our documented 467 plant taxa are utilized in a variety of contexts, including the treatment of different ailments, the management of social relations, the enhancement of hunting skills, the promotion of the development of healthy and strong infants, and the regulation of fertility. We agree with Graham [15] that the classification of plants according to their medicinal or non-medicinal uses is an artificial system designed for illustration and analysis and in no way reflects the traditional hierarchical classification system referred to us by the Cashinahua. It must also be acknowledged that the species reported are only the result of sampling to date and do not represent the complete knowledge of the medicinal plants of the Cashinahua (Table 2).

Table 2 Main therapeutic use categories (≥ 100 UR) presented according to the descending order of the number of UR

The highest number of UR was recorded for socio-cultural uses (398), followed by poisonings (311), infections/infestations (292) and pregnancy/birth disorders (275) as shown in Table 2. The most often quoted health disorders are presented in Table 3, together with the most widespread plant species used to treat them.

Table 3 Ten most frequently reported health problems and species with highest citation frequency for their treatment

The majority of uses in our study represent envenomation, resulting particularly from snake bites as well as stings and bites from other venomous animals, including scorpions and spiders. Snake bite envenomation is an important global public health issue, especially in tropical areas, due to their frequency and resulting morbidity and mortality [61]; globally, an estimated 81,000–138,000 people a year die from snake bites. In 2017, snake bites were re-added to the list of neglected tropical diseases by the World Health Organization [62]. Several recent studies have demonstrated that the most affected group is mainly composed of men working in rural areas, and that snake bites occur mainly during the day, most frequently involve the lower limbs and are mostly caused by the Bothrops genus, which is also the case in South America [63].

Plants for treating snake bites were always the first to be mentioned to us by the research participants and are considered the most culturally important. Poisonings are imminently life-threatening and encounters with venomous snakes in the study area are common. The most common and dangerous bites come from Bothrops spp.—shanu and dunu or “jergón” in Spanish—and Lachesis spp.—kamux or “shushupe”. We were provided with a comprehensive three-stage treatment for snake bite by all of the Cashinahua plant experts we talked to. The plant for the first stage—emergence—is determined (35 species) on the basis of the species of reptile, and always administered externally by squeezing the juice of chewed leaves (or leaves mashed with a few drops of water) into the wound every 3–5 min until the patient evacuates, which is considered the moment when the venom is eliminated. The most important plant for the first stage treatment was considered to be Rosenbergiodendron longiflorum (Ruiz & Pav.) Fagerl—besti bata—that was used for both shanu and kamux, although the types of venom are different. Casearia obovalis Poepp. ex Griseb. and Mascagnia eggersiana (Nied.) W.R.Anderson leaves are frequently used in an emergency to treat different Viperidae bites. Lygodium venustum Sw. is implemented in the treatment of Bothrops bilineatus bites. The leaves of Mascagnia eggersiana (Nied.) W.R.Anderson, Caamembeca spectabilis (DC.) J.F.B. Pastore, Solanum sessile Ruiz & Pav. and Solanum thelopodium Sendtn. are used to treat Lachesis muta bites. The second stage of treatment, that of inflamed wounds, primarily involves bathing the affected part in an herbal decoction (28 species), accompanied by the ingestion of small doses of the same remedy 3–4 times a day, or rubbing the site with leaves pounded in cold water with the same frequency. The final third phase serves to recover the physical strength of the recuperating patient, and can involve 3 species: Dracontium spruceanum (Schott) G.H.Zhu, the corm of which is widely used as a snake bite treatment in the Amazon [64], Cardiospermum halicacabum L. and Solanum mite Ruiz & Pav. Phytochemical screening has revealed that C. halicacabum extract contains glycosides, carbohydrates, flavonoids, phytosterols, phenolic compounds and saponins [65]. During all three phases of treatment, the patient must not move and remain lying in a hammock. According to traditional Cashinahua medicine, the treatment of any venomous snake bite is only successful if strict dietary taboos are obeyed; neither the patient nor his healer must eat cooked cassava throughout the treatment; otherwise, the wound remains inflamed. In case of dietary non-compliance, the inflammation becomes chronic and is treated by rubbing the wound with the leaves of Lepidagathis ipariaensis Wassh. pounded in a small amount of fresh water.

In many parts of the world, scorpion sting envenomation is also a serious health issue that is often overlooked. The true incidence of scorpion sting envenoming is not known because many victims do not seek medical attention. However, it has been estimated that there are approximately 1 million stings per year [66]. Autonomic nervous system mediators released by scorpion venom can cause myocardial damage, cardiac arrhythmias, pulmonary oedema, shock, paralysis, muscle spasms and pancreatitis—all of which can be fatal in young children. In conjunction with intensive care support, early administration of antivenom is highly effective. Nixpu bayai (Piper leucophaeum) was the most widely used species in scorpion bite therapy followed by nidu buxka matsi (Piper marginatum Jacq) and nishi bata (Bunchosia sp.). Slightly grated leaves of the former species are prepared in the form of patarashca, and the warm juice is repeatedly squeezed into the wound.

Of the venomous insect bites, the most widespread are that of the “isula” or buna ant and the Brazilian wandering spider or banana spider (Phoneutria nigriventer)—xina xuku. The isula ant or giant/bullet ant (Paraponera clavata) is a species of hymenopteran insect of the family Formicidae and the only living member of the genus Paraponera. Piper leucophaeum was the most used species in isula and spider bite therapy, along with several Salicaceae taxa such as Casearia obovalis Poepp. ex Griseb. and Lunania parviflora Spruce ex Benth. The most frequently used species to cure venomous spider bites was Aegiphila cuneata Moldenke.

Infections and infestations represented the second most frequently cited category in our study (291 UR), which included different types of herpes, leishmaniosis, conjunctivitis and general infestations. The species with the greatest number of UR in this group were Pseuderanthemum congestum (S.Moore) Wassh. (13), Esenbeckia febrifuga (A.St.-Hil.) A.Juss. ex Mart. (12), Pseuderanthemum lanceolatum (11), Drymonia tenuis (Benth.) J.L.Clark and Piper leucophaeum (10 each). According to our findings, Jacaranda glabra (DC.) Bureau & K.Schum. leaves were used to treat advanced states of leishmaniosis (8 UR). The leaves of Jacaranda copaia (Aubl.) D. Don, as well as those of Piper aduncum, used by the Yanesha displayed favourable activity against Plasmodium falciparum in its chloroquine-resistant strain. Hyptis capitata Jacq. leaves were used for the treatment of skin problems, while another species, namely H. lacustris A. St.-Hil. ex Benth., displayed interesting leishmanicidal activities in the Yanesha pharmacopoeia [67].

Pregnancy, labour and family planning was the third most frequently mentioned category (274 UR), which was cited mostly by female research participants. The species used during gestation, childbirth and the postpartum period were referred to as women’s plants. From this category, Matisia cordata (24 UR), Piper reticulatum (23), Tradescantia zanonia (20), Theobroma cacao L. (19), Pseuderanthemum lanceolatum (18), Quararibea wittii K.Schum. & Ulbr. (17), Acalypha diversifolia Jacq. (10) and Tradescantia zebrina Bosse (9) are most commonly used. With the exception of P. reticulatum, all these plants contain mucilaginous substances (bixtun) and are applied to “increase the phlegm” in the woman’s womb, which is considered conducive to healthy foetal development. These plants are believed to be very useful in facilitating childbirth and for the foetus to grow healthy and strong.

The chopped leaves of various bixtun plants are applied during a bath in the river once a week until the time when labour pains begin. During the first months of pregnancy, a pregnant woman begins to treat herself with Piper reticulatum L.: she drinks a small amount of juice from leaves crushed in a cup of water and rubs her belly with the rest of the grated leaves. A single course every month until the time of delivery ensures that the placenta does not grow too large and that it comes out quickly along with the new-born so that the woman does not suffer from waiting. When contractions begin, the potion made from leaves of P. reticulatum crushed in cold water is taken to accelerate childbirth. Urceolina cyaneosperma (Meerow) Christenh. & Byng and Pavonia fruticosa (Mill.) Fawc. & Rendle are used together to facilitate delivery. Twelve of the documented species are antifertility agents with the predominant species being Chondrodendron tomentosum Ruiz & Pav., Clitoria pozuzoensis J.F.Macbr., Faramea multiflora A.Rich. ex DC. and Rourea amazonica Radlk. Ch. tomentosum is one of the sources of arrow poison curare and it contains, in addition to highly toxic alkaloids, the medicinally valuable alkaloid tubocurarine [68]. To treat women unable to conceive, or to reverse the effect of previously utilized contraceptive plants, the following species, among others, are used: Ruellia proxima Lindau, Pentagonia amazonica (Ducke) L.Andersson & Rova and Urceolina cyaneosperma.

In the treatment of digestive system disorders, the Piperaceae family predominated. In particular, Piper aduncum L., Piper aequale Vahl., Piper costatum C.DC., Piper heterophyllum Ruiz & Pav. and Piper reticulatum L. were frequently used for diarrhoea, vomiting, constipation and as oral hygiene products. Other utilized species included Xylosma tessmannii Sleumer, Uncaria tomentosa (Willd. ex Schult.) DC., Oxalis leptopodes G. Don, Sanchezia oblonga Ruiz & Pav. and Adenocalymma impressum (Rusby) Sandwith. In the context of dental care, it is important to mention the use of several species of Piper involved in the nixpu pima initiation ritual when youngsters (txipax) are introduced to the adult world and for the first time their teeth are blackened with the sap of young shots of nixpu (Piper hispidum Sw. and/or Piper leucophaeum Trel.), which is believed to be the best defence against caries.

Beliefs and symbols in Cashinahua ethnopharmacology

The use of medicinal plants in the study area is highly ritualized, i.e. associated with various kinds of beliefs or with magical or religious practices, as is likely the case in all rural cultures. One important aspect of Cashinahua ethnobotanical practices is the idea that plants play multiple roles at once: medical, social, cultural, pharmacodynamic and symbolic. These roles are subject to significant spatial and temporal variation, as well as frequent interrelations. Ethnobotanical interactions in all of their facets reveal a great deal about broader ecological and social processes [40]. Cashinahua beliefs and practices regarding health, illness and disease cannot be separated from the parallel spirit world, which closely overlays the visible, physical world [15]. The majority of rural cultures do not hold the view that disease and death have an organic or physical cause. The spirit world is the source of both. Kensinger [16] states that one of the ancestral beliefs with which the Cashinahua explain the world refers to the existence of two fundamental aspects: the visible or material side and the invisible side. The visible world is the domain of human beings and all other living things, while the invisible world is the domain of spirits (yuxin) that are impossible to see except in dreams and through hallucinogenic experiences. Informal interviews with healers and their patients revealed a widespread belief in the spiritual and magical origin of diseases. According to different authors [15, 40, 53, 54, 58, 69], the majority of native Amazonian people believe that ancestral wrath, sorcery or an attack by a spirit are the causes of sickness and misfortune. According to Sobiecki [70], they frequently point to strained social relationships. Because specific ailments are perceived only as the result of acts of spiritual agents, the herbalists in our study insisted that we include plants that can be considered “magic” without regard to their status.

Unspecified socio-cultural uses of plants (329 UR) presented in our study were related to enhancing hunting skills, changes in personal odour, behavioural regulation, sexual attraction (“puzanga”), misfortune (“panema”) and sorcery. But the prevalent treatment in this category was to cure symptoms interpreted as a spirit (yuxin) attack. Our findings confirm the statement of Graham [15] that if a yuxin attack were elevated to the rank of illness, it would constitute the most common category of disease reported for the Cashinahua. It would be easy to interpret this as a folk belief, but when we ask what symptoms it manifests, the research participants describe convulsive seizures, often accompanied by loss of consciousness; the affected person is disoriented, suffers great anxiety and cannot control his basic needs. From the point of view of biomedically defined disease states, one might interpret the symptoms of a spirit attack as indicative of some sort of nervous disorder but would never consider a spirit to be an etiologic agent. According to the patient’s behaviour, Cashinahua healers recognize up to six different types of seizures, not unlike epileptic seizures, characterized by their resemblance to animals whose behaviour is similar that of the patient (yawa—peccary, kapa—squirrel, txaxu—deer, isu—spider monkey, xaka—frog, amen—capybara). Each of these kinds of seizure attack is treated with different plants. Graham [15] states that defining and identifying the category of spirit attack is particularly problematic. Treatments in this category include both the use of plants to treat the aftermath of a spirit attack, whose symptoms range from unconsciousness to catatonia to violent range, and the use of plants to protect someone from a spirit attack. In the latter case, plant juice or a decoction is typically taken internally. Warm baths, the application of plant juice to the eye, or both, are typically used as treatment for spirit attacks. Even though mental health issues and psychosomatic disorders have a significant global impact, Western medicine has made little progress in developing long-term treatments that are both sustainable and efficient. To treat these disorders, many traditional societies use ceremonial treatments and mixtures of medicinal plants, frequently to great effect. In order to develop new medications that treat disorders of the nervous system, it may be possible to identify potential future targets through the information gleaned from commonly used traditional treatments [54].

For religious use, only four taxa were presented by research participants, including Banisteriopsis caapi (Spruce ex Griseb.) C.V.Morton bark and the leaves of two Psychotria species, namely P. viridis and P. alba, used together in the preparation of the traditional nixi pae (ayahuasca) entheogenic brew. Another plant species that is added to this vision-inducing potion is Renealmia breviscapa (Poepp. & Endl.) Poepp. & Endl. Alexiades [40] states that ayahuasca is undoubtedly the most widely employed hallucinogen in western Amazonia. Most, if not all, indigenous cultures use hallucinogenic plants as a fundamental part of their healing practices. They frequently play a role that is significantly greater than that of direct therapeutic and calming methods. Research into the tribe’s medicinal plants must therefore include plants used in rituals [46].

New or very rarely reported medicinal plants

Out of 467 plants that were documented, a review of publications on WoS revealed 79 species that have not yet been published as medicinal and have not undergone phytochemical analysis. These species with little or no pharmacological documentation in the scientific literature are distributed among 60 genera and 42 botanical families, with Acanthaceae being the most represented family with seven species, followed by Fabaceae (six), and Araceae and Solanaceae (four each). The fact that they include 172 uses for 79 new or very little-known medicinal plant species is especially remarkable; Table 4 lists the vascular plants among them, arranged alphabetically by species.

Table 4 Medicinal plants used by Cashinahua (Huni Kuin) herbalists in Purus Province, Peruvian Amazon, and previously unreported or very rarely cited for medicinal use or phytochemical analysis

In Table 4, taxa with a frequency of citation (FC) higher than ten are potential candidates for phytochemical or pharmacological research. The following is a discussion of some other notable species in addition to the data presented in Table 2. A wide variety of applications characterize a few species, and five of them stand out because of their adaptability: Aphelandra lasiandra (Mildbr.) McDade & E.A.Tripp., Rourea amazonica (Baker) Radlk., Adiantum poeppigianum C. Presl, Begonia maynensis A. DC. and Leonia glycycarpa Ruiz & Pav.

Of the 79 highlighted species in this research, 11% (12 species) were also described by Manduca et al. [14], who discussed the traditional medicine of the Brazilian Huni Kuĩ (Kaxinawa), with some variations in phonetics and spelling in Hantxa Kuin; 6 species present the same taxonomy, with Leonia glycycarpa Ruiz & Pav., Clavija weberbaueri Mez, Nautilocalyx pallidus (Sprague) Sprague, Aphelandra acrensis Lindau, Cordia nodosa Lam. and Prunus myrtifolia (L.) Urb. being the most frequently mentioned.

Some of the taxa reported in Table 4 have the same uses as other species of the same genus; this is the case for Quararibea wittii K.Schum. & Ulbr. and Q. guianensis Aubl., Drymonia coccinea (Aubl.) Wiehler and Drymonia semicordata (Poepp.) Wiehler, and Paullinia tenera Poepp. & Endl. and Paullinia sp., among others. Similarly, Graham [

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