Food habits related to wild edible plants of the Paraguayan people in the Atlantic Forest

Abstract

The ecological, cultural, and historical complexity of the Atlantic Forest of the Upper Paraná begs a closer look at the local food habits. This is the first study on the current use of wild edible plants by the Paraguayan Mestizo people, who live as migrants in Misiones, Argentina, on the Argentinean–Paraguayan borderland. This study documents 49 wild edible plants, of which wild fruits are the most appreciated food. It finds little connection between eating and healing, and wild edible plants do not appear to be used as medicinal foods. The knowledge of wild food resources is gendered, with men demonstrating greater competence in obtaining wild plants from a greater number of habitat types than women do. These people maintain Guaraní indigenous heritage in preserving the names and uses of numerous wild-grown plants, some of which were first documented in the eighteenth century. In addition, as 86% of the recorded species are native to the region, the Paraguayan people support native biodiversity by using and propagating wild food resources.

Introduction

The food habits of people dwelling in specific environments, including the consumption of wild edible plants, are shaped by various complex social, religious, and economic choices [1, 2–3]. Traditional medical concepts may also play a role in the diet and consumption of wild plants [4, 5]. The ethnobotanical perspective offers insights into the relationship between food habits, or traditional food systems, local agrobiodiversity, environments, and ecologies, which may be valuable for biodiversity conservation and foodways studies [2, 6, 7].

Often, an inquiry into traditional food systems and food habits in a given location contributes to a better understanding of the history and ecology of a given biome [3, 8]. For example, the Atlantic Forest of the Upper Paraná River is blessed with ecological, cultural, and historical complexity, which invites us to look more closely at the local food habits [8, 9, 10–11]. Moreover, the Atlantic Forest biome has undergone intensive changes, especially in the twentieth century [12, 13]; the food choices, particularly those concerning wild-grown edible plants, may tell us about its cultural–ecological entanglements [7, 8, 10, 11].

Wild edible plants are noncultivated. While most plants are native to their natural habitats, this category also includes naturalized and introduced species [1, 14]. Nontended garden escapees, which are usually introduced species, are also considered “wild” in food ethnobotanical studies [1]. Edible wild foods include fruits, starchy tubers, seeds, flowers, and saps; they also include leafy vegetables, i.e., the green parts of plants, such as leaves, stalks, buds, and immature fruits, which are eaten either raw as salads or snacks or cooked [1, 2]. Wild edible plants are regarded as part of agrobiodiversity and constitute a reservoir of genetic diversity suitable for new landraces and varieties. The pool of genes they offer may be of paramount importance because they provide variable responses to environmental stresses, such as changes in soil humidity, drought, fungal diseases, and insects [7, 10, 15].

Studies on the food and medicine continuum noted that numerous edible plants were also employed as medicines in various indigenous contexts among both agrarian and hunter–gatherer communities [4, 5]. Europe has also enjoyed a long tradition of “eat and heal” both in traditional folk systems and in official medical systems, especially in the past [16]. In China, the food and medicine continuum has also played a key role in therapeutic systems since antiquity [17]. However, in lowland South America, researchers have not recorded any such continuum between eating and healing in indigenous traditional cultures, and if so, these were recognized as possible influences of European and Mestizo cultures [18]. Thus, this paper explores the possible relationships between the consumption of wild edible plants and their perceived medicinal value by the Paraguayan Mestizo people, who have been particularly shaped by indigenous Guaraní and European cultures. They were also impacted by the Galenic humoral system during colonial times, whose vestiges can still be found [19, 20–21].

Another question addressed by this paper is whether the knowledge and use of wild edible plants by the Paraguayan Mestizo people are gendered, i.e., whether this knowledge varies between men and women. Our previous study from the same region demonstrated a difference in the knowledge of wild edible plants by Polish migrants and their descendants who arrived at Misiones as part of an organized European migration to the subtropics of Argentina between the end of the nineteenth century and WW2. Men knew significantly more species than women did, which was attributed to men’s wider exploration of diverse environments, principally natural forests, whereas women confined their movements to home gardens, ruderal areas, and agricultural fields, which became their main habitats for collecting wild greens and fruits [11]. Studies examining the distribution of knowledge in a community, such as between men and women, contribute to breaking a vision of gender-neutral ethnobotanical knowledge [22]. Since the role of gender in local systems of knowledge of lowland South America, including the Atlantic Forest, is still relatively poorly studied, the discussion demands new cases.

There has been little research on wild edible plants and their role in the livelihoods and well-being of native and migrant populations in the Atlantic Forest of the Upper Paraná biome. Existing publications concentrate on the Guaraní Indigenous people and European diaspora in Misiones and the Mestizo people, Caiçaras, in the Brazilian part of the Atlantic Forest [7, 9, 10–11, 23]. However, there are excellent historical data from the colonial period on this topic. During that time, the knowledge of Indigenous Guaraní people was first recorded by the Jesuits, whose missions for the Guaraní people were largely located in the Atlantic Forest of the Upper Paraná, in the present central-eastern Paraguay, southeastern Brazil, northeastern Argentina, and part of Uruguay. Indeed, the Misiones region (now the province of Argentina) took its name from the Jesuit missions. The Jesuit masterpiece from that period, Materia Médica Misionera [Materia Medica of Misiones], attributed to Pedro de Montenegro (1663–1726), was devoted to medicinal plants, but some of the included specimens were also edible [24, 25]. This source provides a historical perspective for studying the ongoing use of wild edible plants by the Indigenous and Mestizo people from the region.

There is a lack of scientific publications dedicated to wild foods among the Paraguayan Mestizo people, and to the best of our knowledge, only one brochure on wild edible fruits has been published in Paraguay in recent years [26]. Moreover, a recent comprehensive overview of food studies from the Atlantic Forest biome has included just one publication from the Paraguay concerning the Guaraní people [8]. In view of the above, this paper fills the gap in current documentation of wild edible plants of the Paraguayan people living in one of the richest biomes on the globe [12]. Within this general objective, the paper addresses the following aims: (a) to determine the context of the use of these plants and their role in the diet of the Paraguayan people; (b) to confirm whether a continuum exists between eating and healing, i.e., if wild-grown plants are used as edible medicines or used in prophylaxis; (c) to examine whether the knowledge of these resources is gendered in this society, i.e., if there are differences between men and women; and (d) to compare the richness and pool of the plants used by the Paraguayans with available sources, such as historical colonial ones and those concerning the Guaraní Indigenous people and Polish diaspora living in Misiones, Argentina. This research is important in the context of growing food insecurity, especially in exploring locally available edible flora to mitigate nutrient deficiencies of the local societies. Moreover, it points to potential health benefits that derive from the consumption of wild-grown plants. With this analysis, the paper contributes to food ethnobotanical studies especially to those focused on traditional knowledge in understanding plant biodiversity and local food systems in lowland South America. It also gives bases for more ecological oriented research on the native genetic resources of edible plants and further strategies to protect and promote them as locally healthy food.

Methods

Study area and people

The research was conducted in the Province of Misiones, Argentina, along the eastern bank of the Paraná River, which serves as a border with Paraguay. Both Misiones and eastern Paraguay belong to the same ecoregion and have numerous historical connections. The movement of people across the Paraná River has been frequent since pre-Hispanic times, and until the late nineteenth century, the territory was Paraguayan. Although the Paraguayan Mestizo people are migrants in Argentina, from the perspective of traditional ecological knowledge, their practices should be analyzed as those of local people [20, 27].

The native cover of the study area is semideciduous Atlantic Forest. The climate is subtropical and humid, with a mean annual rainfall of 2000 mm and a mean annual temperature of 20 °C. The Atlantic Forest originally covered approximately 1,500,000 km² [12]. At present, nearly 40% of its original cover has been preserved in Misiones, Argentina, in contrast with Brazil and Paraguay, each of which retains only 8% of the original forest preserved mainly in protected areas [12, 13]. Despite the large anthropogenic disturbances, this ecoregion still remains among the most biodiverse biomes in the world [8, 12].

In precolonial times, this region was inhabited by different Guaraní groups: mobile Indigenous horticulturists [23]. During colonial times, until the late eighteenth century, Misiones was dotted with Jesuit missions, where the Guaraní lived. The Jesuits introduced herbs and crops of Mediterranean and Asian origin, which they propagated among the Guaraní and they also studied plants used by this Indigenous group [24, 28].

For its part, Paraguay has been isolated politically and geographically throughout its history. The colonization process in Paraguay was based on settlement and farming rather than looting minerals; this led to the early emergence of an entirely new Paraguayan bilingual society that was vastly influenced by Guaraní heritage but also impacted by Spaniard culture [19]. In the nineteenth century, owing to the War of Triple Alliance (1865–1870), in which Paraguay fought against three allied aggressors, Argentina, Brazil, and Uruguay, the country was left devastated. Paraguay also lost Misiones to Argentina, with the border being settled on the Paraná River [27]. The authorities of Argentina wanted to prevent the influx of Paraguayans from the east to the newly acquired territory of Misiones. They were also in favor of the European colonization of Misiones. As a consequence, massive waves of European migrants arrived at Misiones between the end of the nineteenth century and WWII. Even so, the long Argentina–Paraguay border on the Paraná River (1699 km) has been porous, leading to the continuous movement of people and goods [27]. The latest census, from 2022, estimated that 522,598 Paraguayan migrants are living in Argentina, constituting 27% of all migrants therein and forming the largest Paraguayan collective outside the country [29]. Since the mid-twentieth century, the main reasons for migration have been political and economic, such as finding better access to public health and social security systems in Argentina [20].

Data collection

The fieldwork for this study was conducted in four localities on a spectrum from rural to urban areas along the Paraná River. They are the following: Wanda, with Puerto Wanda as one unit (25° 58′ S, 54° 34′ W); Piray Km 18 and Puerto Piray, which belong to the same municipality (26° 28′ S, 54° 43′ W); and Puerto Leoni (26° 59′ S, 55° 09′ W) (Fig. 1). The data were collected in 2014, 2015, and 2019.

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Fig. 1

Map of study localities in the Province of Misiones in Argentina, South America

Sixty people participated in the study, of whom 44 were Paraguayan migrants originating from eastern Paraguay, within the Atlantic Forest ecoregion, and 16 were descendants from the first generation born in Argentina. The group included 32 women (mean age 62.8, min. 33, max. 95) and 28 men (mean age 61, min. 35, max. 82). The majority were bilingual in Guaraní and Spanish. All operated small farms, working on 1–2 hectare agriculture plots, cultivating staple crops, such as maize (Zea mays) and cassava (Manihot esculenta), or just maintaining a home garden with separate units for vegetables and herbs, the latter being the case in towns (Wanda and Puerto Leoni). Men also find employment in the forest industry, whereas women usually remain housewives or find informal employment as domestic helps. The snowball technique [2, 3] was used to contact the study participant, with the inclusion criteria as following: being adult Paraguayan migrant hailing from the same biome of the Atlantic Forest or the 1st generation descendants who identify themselves with the Paraguayan culture and country; showing interest in plants (food and medicinal), being involved in agriculture or having agricultural background. During sampling, the author’s attempt was to maintain a similar number of women and men for further comparative analysis.

Information on wild plants was obtained via the free listing technique [30], complemented with walks in the forest and other environments where the study participants usually collected their wild-grown plants. Additionally, open-ended conversations and observation were implemented. The research opened up with an oral free listing task during which the interviewees were asked to elicit all wild edible plants they knew and had previously consumed. There weren’t any restrictions concerning the time needed for producing a list or the number of listed items. When the person finished listing the known and consumed plants, additional questions were posed to each elicited plant species regarding the following: (a) the part of the plant used; (b) ways of preparation and consumption; (c) the methods of acquisition and plant availability; (d) the taste appreciation score (from 1 to 10); and (e) the frequency of consumption and the last time the person ate a given plant.

Walks in the forest were performed to collect voucher specimens with the study participants. These walks were not a systematic plot or transect sampling, but rather they were guided by the study participants to the forest sectors which they knew and where they collected their edible plants. During these walks, only photo documentation was done (without any recording) and notes were taken on spots, which included not only plant names and their uses, but also chemosensory cues, such as smell, taste, and texture, and also some ecological cues about specific habitats they grew in. To better contextualize the use and consumption of wild-grown plants, open-ended conversations were performed with volunteering men and women from the same pool of study participants. Moreover, to understand the place of wild plants in the lives of the Paraguayan people, the participants were observed during plant gathering, preparation, and consumption, which were also documented with photos. The documented botanical species were supported with herbarium specimens stored in the CTES Herbarium at the Instituto de Botánica del Nordeste (IBONE), Corrientes, Argentina. The botanical identifications were largely done by the author, but some experts working with particular botanical families in this herbarium participated in the identification of certain specimens, and their names were included in the plant labels. These were: Sara G. Tressens for Myrtaceae, Maximiliano Dematteis for Asteraceae, Ricardo Vanne for Fabaceae botanical families. Moreover, the herbarium expert technicians were involved in this process (for both medicinal and edible plants): Gelina Pieszko and Walter Medina.

Analyses

The plant names were checked and confirmed via Plants of the World Online (POWO) [31]. The origin of the plants (native or introduced) was established through the Catalogue of Vascular Plants of the Southern Cone [32]. The frequency of mentions was calculated to act as a proxy for the cultural importance of particular wild edible plants. Another similar study demonstrated that other cultural indices may be redundant in the context of edible plants in the Neotropics: the salience index provides similar information to the frequency of mentions, and the cultural food significance index (CFSI) may be biased toward certain food types [11]. However, a taste appreciation score was calculated as the mean value for each plant; this value was presented in the Results for the ten most frequently mentioned plants as an additional indicator influencing the frequency of use. The significance of the differences between men and women was calculated via the Mann–Whitney U test. Statistics were calculated using the open access Python library (SciPy).

The recorded species were compared with data given in other studies: these included a recent publication of wild edible plants used by Polish migrants and their descendants in Misiones, Argentina [11], two studies on edible plants by the Guaraní people [9, 23], and one colonial Jesuit volume: Materia Medica Misionera [24]. Plants used as additives to mate (Ilex paraguariensis infusion) and tobacco were excluded from the list of Guaraní people [23], because they may also have medicinal uses. The degree of similarity between the wild edible plants used by the Paraguayan migrants and those used by the Polish diaspora, the Paraguayans, and the contemporary Guaraní people was estimated on the basis of the Sørensen similarity coefficient [33]. The Sørensen index is calculated by doubling the number of elements common to both sets, dividing this value by the sum of the number of elements in each set, and multiplying the result by 100 to obtain a percentage value.

Results

Characteristics of wild edible plants used by the Paraguayan people in Misiones

In total, 49 botanical species corresponding to 47 ethnotaxa were recorded: the ethnotaxon “guayaba” was found to comprise three different species. These species belong to 25 botanical families, of which the most representative is the Myrtaceae (n = 13; 26.5% of all species), followed by the Arecaceae (n = 3) and Asteraceae (n = 3); the remainder of the botanical families are represented by two or one species (Supplementary Table 1). From the pool of 49 recorded plants, 42 species (86%) were native to the Atlantic Forest, one species was naturalized (Citrus aurantium), and six had been introduced to the region: three of them are fruit trees, such as Eriobotrya japonica, Hovenia dulcis and Morus alba, which were already known to the study participants from Paraguay and remaining three were herbal species brought by the Jesuits and European migrants (Lactuca virosa, Nasturtium officinale, Sonchus oleraceus).

Documented plants occur in a wide range of habitats, while trees and palms predominate in the forest (first growth disturbed forest, second growth forest), often along streams, many of them, particularly from the Myrtaceae family and two palms: Acrocomia aculeata and Syagrus romanzoffiana are also found in the forest edge, along the pathways, in ruderal areas of early and middle ecological succession (locally called capuera). Wild-grown trees and shrubs bearing edible fruits are often protected in the agricultural fields (seeds dispersed by birds and mammals are left to grow). Sometimes young seedlings or young plants are transplanted to home gardens, or they are sown directly from seeds in the desired parts of the agricultural lands or home gardens. This usually happens with highly praised native Myrtaceae, Annonaceae, and Fabaceae fruit species. However, some trees are associated with forest environment and very rarely transplanted to home gardens, such as Araucaria angustifolia, Celtis iguanaea, Chloroluma gonocarpa, and Jacaratia spinosa. The practice of transplanting and sowing of wild fruit-bearing trees close to the domestic areas has been practiced for two main reasons: first, to have these resources at hand; and secondly, to preserve them from disappearance. The interlocutors, especially those living surrounded by monocultures of introduced pine and eucalyptus forests, were concerned with the little availability of native fruit-bearing trees in their environment. On the other hand, they felt constrained by the limited space (1–2 ha plot or a home garden), and therefore, they had to make choices about which trees and shrubs to plant and protect in their home gardens. In these circumstances, multiple functions of plants, such as food, medicine, and shade, increased their chances to be protected, something I will return to in the discussion. Edible leafy weeds which are considered useful as additives to salads or other side dishes are not eradicated from home gardens but rather tolerated. They are also obtained from ruderal areas and agricultural fields.

The ten most frequent plant species were mentioned by more than 50% of the study participants (> 30) (Fig. 2). All are native trees bearing edible fruits and can be considered the core wild edible plants known and used by the Paraguayan Mestizo people living in the Atlantic Forest region. While the most frequently consumed species are the palm coco (Acrocomia acuelata), the Myrtaceae tree “yva hai” (Campomanesia xanthocarpa), and the forest epiphyte “guembe” (Philodendron bipinnatifidum), the most appreciated plant, owing to the taste of its fruits, is “ñangapiry” (Eugenia uniflora), followed by “pakuri” (Garcinia brasiliensis) and “aratiku” (Annona neosalicifolia) (Fig. 3).

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Fig. 2

Plant species most frequently consumed and appreciated for their taste by the study group

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Fig. 3

Examples of frequently mentioned wild edible plants by the Paraguayan Mestizo people, Misiones, Argentina. A Ripening fruits (nuts) of “coco”-Acrocomia aculeata (Arecaceae) (50 mentions), Puerto Leoni; B Unripe fruit of “yva hai”-Eugenia pyriformis (Myrtaceae) (30 m.), Puerto Wanda; C Unripe fruits of “guembe”-Philodendron bipinnatifidum (Araceae) (48 m.), Puerto Leoni; D Ripe fruit of “ñangapiry”-Eugenia uniflora (Myrtaceae) (46 m.), Puerto Leoni

The most frequently consumed plant parts are fruits. Most are eaten fresh as snacks, straight from trees, palms, or bushes without any further processing (31 out of 49 recorded species) (Table 1). For example, Myrtaceae trees and shrubs bear fruit once a year during the spring and early summer (October–December), depending on the species, so they are enjoyed during a relatively short period of time (they may have a second less abundant fruiting season in the fall: May–June). These plants are consumed on hot spring days during family picnics at the streams and by children during school breaks, and they give an excuse for a break for those working in the forest. Hence, there is a conjunction between the consumption context (good weather, picnic, school or work break) and the relatively short period of foraging, which strengthens the valuation of these plants and their tastes. A few fruits are processed occasionally into marmalades (Psidium spp., Campomanesia xanthocarpa), compotes (Eriobotrya japonica) or juices (Citrus aurantium, Passiflora alata), and are consumed in salads as vegetables (Melothria cucumis) or roasted on embers or in an oven (Bromelia balanasae, Jacaratia spinosa) in order to eliminate their spiciness (hotness).

Table 1. Categories of use and the corresponding wild edible species

Category of use	Local name(s), Latin name, plant part used and mode of preparation
Additives to other dishes	coco (Acrocomia aculeata), ground seeds are added to yerba mate drink or taken with hot milk in a mate gourd; apepu (Citrus aurantium) fruit juice added to lemonade, salads, tereré (cold yerba mate infusion); palmito* (Euterpe edulis) palm heart fried, additive to main dishes, "it is another cabbage"; ovenia (Hovenia dulcis) grated seeds are used as a substitute for coffee; ortiga grande, pyno (Urera baccifera) condiment and additive to food; ortiga chica (Urtica circularis) condiment and additive to food
Desserts, marmalades, compotes	guavira* (Campomanesia xanthocarpa) fruits in marmalade; níspero* (Eriobotrya japonica) fruits in compote; pitanga (Eugenia uniflora) dessert fruits; tuna (Opuntia arechavaletae) raw fruits after peeling as dessert; mburukuya (Passiflora alata) added to desserts; yva puru, yaboticaba (Plinia cauliflora) dessert fruits; arasa*, guayaba* (Psidium guajava, Psidium guineense, Psidium cf. australe) dessert fruits and in marmalade;
Raw fruit snacks	kokû (Allophylus edulis); araticu'ñu (Annona emarginata); araticu (Annona neosalicifolia); karaguata (Bromelia balansae); ñanadu apysa (Campomanesia guaviroba); ñanadu apysa ñu (Campomanesia guazumifolia); guavirami (Campomanesia sp.); guavira (Campomanesia xanthocarpa); tala (Celtis iguanaea); aguay (Chloroluma gonocarpa); níspero (Eriobotrya japonica); yva mbo py, cerella (Eugenia involucrata.); yva hai (Eugenia pyriformis); ñangapiry, pitanga (Eugenia uniflora); guapo'y, higuera (Ficus spp.); pakuri (Garcinia brasiliensis); ovenia (Hovenia dulcis); inga (Inga marginata); ingá guasu (Inga vera); mora (Morus alba); guaviju (Myrcianthes pungens); guembe (Philodendron bipinnatifidum); guaporaity (Plinia rivularis); mora rosada del monte (Rubus rosifolius); zarzamora (Rubus urticifolius); espina colorada (Solanum sisymbriifolium); pindó (Syagrus romanzoffiana)
Roasted food	coco (Acrocomia aculeata) roasted seeds eaten as a snack; pino paraná (Araucaria angustifolia) roasted seeds eaten as a snack; yacaratia (Jacaratia spinosa) fruits roasted on embers and eaten
Fresh leafy vegetables	tacuará (Guadua spp.) young bamboo shoots; palmito (Euterpe edulis) palm heart eaten raw in salads; achicoria silvestre (Hypochaeris chillensis) raw leaves in salads; lechuga japonesa (Lactuca virosa) raw leaves used in salads; mintruz (Lepidium auriculatum) raw leaves used in salads; pepinillo (Melothria cucumis) raw fruits used in salads; berro de agua (Nasturtium officinale) raw leaves used in salads; makysi, trebol (Oxalis debilis) raw leaves and bulbs added to salads or eaten as snacks; llantén (Plantago australis) raw leaves added to salads; lengua del buey (Sonchus oleraceus) raw leaves added to salads; pindó (Syagrus romanzoffiana) palm heart added to salads;

The categories in which plants are consumed fresh are “raw fruit snack” and “fresh leafy vegetables”; the category “roasted food” marks a basic process of roasting to enhance the taste or to improve the edibility; the remaining two categories: “additives to other dishes” and “desserts, marmalades, compotes” include both freshly consumed, cooked, and fried food, but the one which requires heat processing is marked with (*)

The Paraguayan Mestizo people observe some proscriptions regarding mixing certain foods. They are particularly concerned about fruits. For example, fruits should not be eaten after main dishes or before or after hot and cold infusions of yerba mate (Ilex paraguariensis). In fact, they should be eaten on their own, preferably not mixed (only one species at a time), and there should be a sufficient digestive span between eating fruit and drinking water, yerba mate, and eating dishes such as meat or hot pots.

The second most frequently used plant part is the leaf, with a score of eight species out of 49. Apart from Hypochaeris chillensis, a native herbaceous plant mentioned in Materia Medica Misionera [24], leafy vegetables are not popular in Paraguayan society. The interlocutors mentioned that they did not enjoy eating green salads and that it was a food of the “colonos”- European migrants and their descendants. When salads are prepared, they are typically cultivated cabbage (Brassica oleracea) or lettuce (Lactuca sativa). When asked about the food they used to eat in the countryside in Paraguay, where they grew up, most people indicated what was produced on their farms: landraces and varieties of maize, cassava and rice; batatas; beans, including lentils, peanuts, pumpkins, and chives; and livestock animals and their derivatives. During fieldwork, I frequently observed people eating one-pot stews with meat and reviro: wheat flour fried with lard; practically no greens were added. Moreover, the cultivation of delicate lettuce, rocket, spinach, tomatoes, cucumbers, and other salad vegetables is complicated by difficulties in watering gardens, and these vegetables are considered food habits of “colonos” as well.

A few seeds rich in oils are also consumed, of which the most popular is “coco,” Acrocomia acuelata, which is considered a very powerful food that is eaten raw and roasted. Many interlocutors could also remember that when they were children, they used to have grated or crushed seeds of “coco” with hot milk sipped from the gourd, a preparation similar to yerba mate hot infusion, and it was served for breakfast. The interlocutors emphasized that at that time of their childhood, everyone had a cow in the Paraguayan countryside, and milk was drunk on a daily basis. An interesting adaptation was the use of grated seeds of the introduced tree species Hovenia dulcis as a substitute for coffee. Finally, three species are used for their soft parts, namely Euterpe edulis and Syagrus romanzoffiana for their palm hearts and Guadua sp. for young bamboo shoots, which are consumed fresh or prepared into preserves.

Food medicine continuum?

Wild edible fruits are perceived as healthy food resources by the Paraguayan people in Misiones, but they are not connected with any particular medicinal property. Fruits are eaten mainly as raw snacks because they have an interesting taste and scent that are different from those of cultivated plants. Additionally, their tastes are blended with memories and nostalgia of Paraguay. The only fruit that appears to have medicinal value is the juice of bitter orange, Citrus aurantium, which is used to treat respiratory illnesses. Wild leafy vegetables are considered healthy foods, especially Hypochaeris chillensis, Sonchus oleraceus, Lepidium auriculatum Nasturtium officinale, and Plantago australis. They are used, however, by a small group of people who follow healthy trends or are restricted to diets owing to diabetes or high levels of cholesterol. In particular, the bitter taste of Hypochaeris chillensis, Sonchus oleraceus and Lactuca virosa is viewed as an anti-fat and anti-sugar agent capable of “burning” sugars and fats. Two nettle species, Urera baccifera and Urtica circularis, are consumed as healthy additives to salads. It seems that medicinal foods are not common in the Paraguayan Mestizo culture and that the domains of food and medicine are separate. However, it is important to note that 28 out of 49 recorded species (57%) are also used medicinally, but different plant parts are used as food and medicine.

Gendered knowledge

The study participants reported, on average, 13.2 ethnospecies. The mean number of ethnospecies mentioned by men was greater than that mentioned by women, and this difference was statistically significant. Table 2 compares the results from this study with a similar pool of data from the Polish diaspora in the same region [11]. Both groups demonstrated similar patterns in terms of knowledge of wild edible plants. It appears that the predilection for wild-grown fruits is associated with their growth in the forest and greater exploration of these habitats by men than by women. Leafy vegetables can be more easily found in the ruderal areas around homes or even in home gardens, which are usually tended by women, but this type of wild food is not frequently consumed.

Table 2. Familiarity with wild foods among men and women from the Paraguayan and Polish diaspora in Misiones

Study group	Mean number of species	Mean number of species (men)	Mean number of species (women)	Significance (men vs. women)
Paraguayan diaspora	13.2 ± 4.85 SD (median-12)	14.4 ± 4.02 SD (median-14)	12.16 ± 5.44 SD (median-11)	p = 0.0170
Polish diaspora	13.3 ± 4.7 SD (median-13)	15.3 ± 4.4 SD (median-16)	11.5 ± 4.3 SD (median-11)	p = 0.0068

Continuation and change in the migration process

No changes in the plant name used were observed between the surveyed migrant descendants of the Paraguayan people (n = 44) from the first generation born in Argentina (n = 16): both groups preserved the Guaraní language and the names of the plants. The mean number of known plant species in the Paraguayan migrant group (n = 13.36 per person) was only slightly greater than that in the other groups (n = 12.75 per person). However, the migrants were more likely to mention the names of wild edible plants that they had known and consumed in Paraguay but that were not easy to find in Misiones, such as “guaviju” (Myrcianthes pungens) and “pakuri” (Garcinia brasiliensis), “inga guasu” (Inga vera), “yacaratia” (Jacaratia spinosa), “araticu riu” and “chirimoya del campo” (not identified). Some others managed to transplant them from Paraguay to their home gardens or agricultural fields in Misiones.

Comparison with other ethnic groups from the Atlantic Forest of the Upper Paraná River

The Paraguayan participants shared 76.4% similarity with the Polish diaspora from Misiones, indicating that both recognized a corpus of wild edible plants found in the region, preserving some ethnic preferences and particularities (Supplementary Table 1). For example, while the Paraguayan people extensively explore wild-grown Myrtaceae tree species, the Polish population is more likely to exploit wild green vegetables.

The Paraguayans also demonstrated 58% of similarity with the contemporary Guarani Indigenous people from Misiones, according to the Sørensen coefficient. The Guarani people preserve a wider scope of uses for wild edible plants and their forms of consumption. While a record of ethnobotanical knowledge among the Guaraní in Misiones in the mid-twentieth century by Martinez-Crovetto mentioned that the Guaraní used 60 botanical taxa, four of these could not be found in any indexed botanical taxonomic database; in addition, Celtis pubescens is a synonym of Celtis iguanaea and Eugenia uvalha, a synonym of E. pyriformis, Psidium australe and P. cuneatun; hence, the updated botanical richness in Martinez-Crovetto’s work is 53 species. This Guaraní source from the mid-twentieth century shares 55.4% similarity with the contemporary Paraguayan Mestizo community in Misiones. Finally, 12 edible plants recorded in Materia Medica Misionera were also documented in the present study. Nine of them are edible fruits, one is an edible tuber or rhizome (Oxalis debilis), and two are wild leafy greens: Hypochaeris chillensis and Plantago australis; however, the latter two were only used in a medicinal context during the Jesuit times [24].

Discussion

Wild edible plants, their richness, and significance to Paraguayan people living in Misiones

The number of plant species, i.e., the ecological richness, identified in this study is lower than that observed for Indigenous Guaraní groups of Misiones but higher than that of European (Polish) migrants and their descendants from the same region. The documented botanical richness of wild edible plants among the Paraguayans is within the range recorded among other native groups from Argentina and the South Cone of South America. For example, the Chorote people from the Gran Chaco (northwest Argentina and southwest Paraguay), previous hunter–gatherers, use 57 wild edible plants in a diverse way [6]. In central Argentina, 45 species of wild edible plants were documented among the rural population of mixed origin [34]. In another arid region of Argentina, Patagonia, the Mapuche Indigenous group reported a knowledge of 42 species, but the authors noted that this knowledge of wild food had been eroded in this population [35]. In the rural communities of the Brazilian Pantanal, along the Paraguay River, Bortolotto et al. [36] reported 54 wild-grown species with edible uses. Interestingly, as noted in the present study, Acrocomia acuelata was also found to be the most important and versatile species along the Paraguay River in Brazil [36].

On the basis of this short overview, with 49 botanical species, the Paraguayan Mestizo people can be considered just as knowledgeable regarding wild edible flora as other populations from the South Cone of South America. However, these resources contribute little to the diet of Paraguayan individuals as components of daily and festive dishes: the intake of wild edible plants comes down to seasonal snacking on fruits and nuts, and the exploration of leafy greens is limited to a few species used by a small group of people. Very little information about the consumption of wild leafy vegetables was noted in the historical sources and those concerning the Guaraní people from Misiones, and the Caiçaras, people of mixed Indigenous and European origin, living in the Atlantic Forest of Brazil [9, 10, 23]. The preference for wild fruits over wild greens can be attributed to cultural and ecological traits: the traditional pattern of food habits in the region and the strong prevalence of fruits among the edible wild flora in the Neotropics [7, 8, 9–10].

In addition, little connection was observed between eating and healing in the studied group, i.e., wild edible plants are not part of local strategy to supplement micronutrients. But as Johns suggested: “people eat food not nutrients” [37]. In the context of Paraguayan people, this means that there exist mechanisms for ingestion of secondary compounds, such as extensive knowledge and efforts to protect these resources to increase their availability, but they may not be explicit. During fieldwork, I noticed that people paid more attention to daily use of medicinal plants in their prophylaxis or treatment than to wild edible foods. Indeed, the Paraguayan Mestizo society has been reported to demonstrate an extensive and constantly adapting understanding and use of medicinal plants [20]. In this society, the intake of micronutrients and medicinal compounds is mainly regulated by the habit of adding medicinal plants to hot and cold yerba mate infusions (Ilex paraguariensis). A previous study on the Paraguayan Mestizo people living in Misiones documented close to 100 medicinal species taken as infusions with yerba mate on a daily basis as prophylaxis or for the treatment of particular health problems, including chronic diseases [38]. Another reason for little interest in wild edible plants apart from seasonal snacking is the farmer-style high caloric diet, which is rich in carbohydrates (cassava, maize, batatas, plantains, rice, wheat flour, beans), and fats. As such, the most significant plant among the wild edible plants is Acrocomia acuelata, which produces oily fruits. There is little space for leafy greens and salads in the Paraguayan diet [21]. These food habits are connected to humoral concepts, which impose some restrictions on the appropriate time for the ingestion of fruits [21]. Moreover, the interlocutors frequently stated that fruits did not combine very well with drinking yerba mate (neither before nor after), which is taken at least twice a day and is the most important social drink of the Paraguayan people par excellence [38]. Nonetheless, the knowledge of wild edible plants is still rich in this society and transmitted from generation to generation; therefore, if needed these people have the capacity to switch to wild foods. If people’s knowledge and the resources themselves were soundly promoted for counteracting nutritional deficiencies, the Paraguayan people would most probably include them into their diet with higher frequencies.

The high similarity in the corpus of wild foods used by the Paraguayan Mestizo people and the Polish migrants confirms our previous findings that these were the Paraguayan people who played a vital role in familiarizing the European newcomers with native plants of Misiones [11, 20]. The Paraguayan people appear to be a “bridging” ethnic group in Misiones. On the one hand, they share many names and uses of wild edible plants with the Guaraní people, although the latter exhibit greater knowledge of the richness and diversity of wild edible fruits [9, 23]. On the other hand, their diet shares some wild greens with that of the European migrants and their descendants, although the Europeans demonstrated a greater richness and diversity of their use, especially as fresh leafy vegetables.

Native biodiversity of wild edible plants

Notably, the Paraguayan people support native biodiversity by using and propagating wild edible plants. The great majority (86%) of recorded species are native to the region, and they are facilitated by human management and practice. Compared with populations of mixed origin in the South Cone in South America [10, 11], the Paraguayan Mestizo people rely almost exclusively on native plant species as wild foods. Through their management, the Paraguayan people from the Atlantic Forest contribute to a wider distribution, as indicated by the greater range of habitats and the greater abundance of native useful flora. A similar phenomenon was observed among the Caiçaras in the Brazilian part of the Atlantic Forest, although not on such a scale [10]. Twenty-five perennial species bearing edible fruits were also found to be managed in home gardens by people of mixed origin in Puerto Iguazú in Misiones, a border town with Brazil and Paraguay. The most common management techniques used for those plants are sowing, tolerating, and transplanting, especially in home gardens adjacent to forest areas [39]. However, the efforts of local people in Misiones based on transplanting, protecting, and cultivating wild edible plants are not sufficient strategies to preserve these resources. They should be coupled with the government strategies to protect the habitats, by introducing more regulations to prevent the replacement of natural forests with pine and eucalyptus monoculture. Moreover, it is important to let the local people know that their knowledge is a marker of resilience, which mitigates the processes of food industrialization. This can be done, for example, by creating more opportunities to sell the local foods on the local markets, called ferias francas.

The Atlantic Forest of the Upper Paraná Basin is characterized by a great diversity of Myrtaceae trees and shrubs [32]. The outstanding richness and frequency of use of Myrtaceae plants recorded herein are in line with previous studies conducted among the Guaraní people [9, 23]. They were also most frequently mentioned among fruit trees in Materia Medica Misionera [24, 25]. The Myrtaceae family also stood out in ethnobotanical studies from the Brazilian part of the Atlantic Forest [7, 10], as well as in the recently published overview of edible plants from the Atlantic Forest biome [8]. This may be due to the conspicuous features of these plants: their edible fruits, the medicinal properties of their leaves and bark, their pleasant fragrance, their relatively easy propagation from seeds coupled with their abundance in the Atlantic Forest [39]. Thus, local populations actively participate in the incipient domestication of Myrtaceae plants bearing edible fruit, altering their genetic structure and their abundance and distribution [8, 10, 39]. Moreover, the names of the Myrtaceae ethnospecies were largely preserved in the Guaraní language from the colonial period and were transmitted to the European migrants in the twentieth century [11]. Therein, knowledge about Myrtaceae shrubs and trees is an example of persistent traditional ecological knowledge, which has not only been retained by the Indigenous and Mestizo societies but also transmitted to new European settlers. This preservation of knowledge and related practices must have influenced both the use and the management of these species.

The fruits of the Myrtaceae have a complex taste: both sweet and sour, with slight astringency, and the degree of sourness and astringency differs from species to species. As some of these fruits contain tannins (Campomanesia xanthocarpa, Eugenia uniflora) or saponins (Psidium spp.), which make them astringent [40], the Paraguayan people ingest plant micronutrients by consuming secondary compounds from wild edible fruits; however, this is not a conscious process, as is the case for the ingestion of medicinal plants with yerba mate infusions. Nonetheless, the multi-contextual ingestion of fruits as food and of leaves or bark in medicinal infusions increases the potential of exposing Paraguayan people to active substances found in the Myrtaceae species.

Conclusions

This is the first study on the current use of wild edible plants by the Paraguayan Mestizo people, who live as migrants in Misiones, Argentina, on the Argentinean–Paraguayan borderland. The Atlantic Forest in Misiones is better preserved than that in eastern Paraguay; as such, the Paraguayan people were able to continue their traditional ecological knowledge in the realm of wild edible plants. However, they also suggested that the progressive expansion of pine forest monocultures might reduce the availability of the native trees and palms. These people maintain Guaraní heritage in preserving the names and uses of numerous wild-grown plants, some of which were documented in the eighteenth century. They also served as experts to migrating European settlers in familiarizing them with the edible native flora of the Atlantic Forest of the Upper Paraná River.

The botanical richness documented in this study (49 species) indicates that the Paraguayan people are knowledgeable about wild edible plants. Additionally, men demonstrate greater expertise in the identification and exploration of these resources in the forest. However, wild edible plants do not play an important role in their daily diet or as festive food; their consumption is mainly maintained by snacking on seasonal fruits, which have appealing tastes and scents. Wild leafy vegetables are appreciated by a small group of people. Wild edible plants are not an explicitly important source of micronutrients or food medicines, as Paraguayans regulate these needs through the daily intake of medicinal plants together with their cherished infusions of yerba mate. Therefore, a large amount of knowledge of wild edible plants stems from the extensive, more general, traditional ecological knowledge of these people and from the richness of edible flora in the Atlantic Forest rather than from their deep interest in this type of food and conscious strategies in using these resources. Owing to their strong reliance on native plants, the Paraguayan Mestizo people promote the diversification of the native flora, especially the fruit-bearing trees and shrubs of the Myrtaceae family, through the active management of native plants.

Acknowledgements

Special thanks are given to the Paraguayan migrants living along the Paraná River in Provincia de Misiones, Argentina, for their generosity in sharing their traditional ecological knowledge and for their confidence in the research project. I would like to thank Héctor Keller, the Herbarium staff in IBONE Corrientes, particularly to Gelina Pieszko and Walter Medina, for their help in the exact identification of the herbarium specimens. I am also grateful to Bernard Kujawski for his help with the Python library and for generating Fig. 2.

Author contributions

MK carried out the fieldwork, filled the database, performed the analysis, wrote the draft of the manuscript and its final version.

Funding

The research was financed by funds from the National Science Centre, Poland, research grant nos. 2018/31/B/HS3/03019.

Availability of data and materials

All the relevant data from the research on wild edible plants among the Paraguayan Mestizo people living in Misiones, Argentina, were included in this paper in a form of descriptions, tables, figures, and references to herbarium collection. Moreover, an extensive database was attached as supplementary data (Supplementary Table 1). Other field materials such as field diaries cannot be shared openly, because they contain sensitive information about the study participants. The funding agency, National Science Centre in Poland, did not require for this project to place the field data in any public repository.

Declarations

Ethics approval and consent to participate

Oral informed consent was obtained from each adult Paraguayan person and their descendants who volunteered to participate in the study. Consent to publish any photograph included in this research has been obtained from the study participants. The study followed the ethical protocols established by the AAA and the International Society of Ethnobiology. The project did not require approval from the ethical committees of University of Łódź, where the project was implemented or from Argentinean institutions. The necessary permits were obtained from the Ministry of Ecology in Posadas, Argentina, and deposited with herbarium specimens in IBONE, Corrientes, Argentina. No other permits were needed, as the study took place outside protected areas.

Consent for publication

The author of the manuscript is accountable for all aspects of the accuracy and integrity of the manuscript in accordance with ICMJE criteria. The article is original, has not already been published in a journal, and is not currently under consideration by another journal. The author agrees to the terms of the BioMed Central Copyright and License Agreement.

Competing interests

The author declares that she has no known competing financial interest or personal relationships that could have appeared to influence the work reported in this paper.

Publisher's Note

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

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