Wild and native plants and mushrooms sold in the open-air markets of south-eastern Poland

Background The study of plants and fungi sold in open-air markets is an important part of ethnobotanical enquiry. Only few such studies were carried out in Europe. Methods Four of the largest open-air markets of south-eastern Poland were visited regularly, and the plants sold in them were recorded between 2013 and 2015. The aim of the study was to record native and/or wild species sold in the markets. All the plants sold in the markets were photographed regularly. In each market, 25 sellers were interviewed. Voucher specimens were collected and fungi were identified using DNA barcoding. Results Altogether, 468 species of plants were recorded, 117 of them native to south-eastern Poland – 19 only collected from the wild and 11 both wild and cultivated. Seventeen of the species are under legal protection. Most protected plants were sold from cultivation, although proper authorization procedures had not been performed. Thirty-two species of fungi were sold (including two cultivated species), all of them for culinary purposes. Two species (Lactarius quieticolor, Leccinum schistophilum) are new to the mycobiota of Poland. Ornamental plants constituted a large section of the market, and they dominated the group of native species. Food plants dominated among wild-collected plants and were sold mainly as fruits for jams, juices and alcoholic drinks, or as culinary herbs. Very few medicinal or green vegetable plants were sold. An interesting feature of the markets was the sale of Ledum palustre as an insect repellent. Conclusions Finding two species of fungi which are new to Poland highlights the importance of DNA barcoding in ethnomycological studies. Most items in the markets are ornamental plants, or edible fruits and mushrooms. Very few medicinal plants and green vegetables are sold, which differentiates the markets from southern European ones. Such a pattern is probably the model for most central European markets.


Background
The study of plants sold in open-air markets is an important part of ethnobotanical enquiry [1,2]. Plants which are sold in such places are usually those which are culturally the most salient. In traditional agricultural societies the market is often the main source of goods sold and bought by villagers. Even with the advent of regular shops and supermarkets in more modernized societies, open-air markets remain an important centre of plant commerce for both urban and rural dwellers. Many of the plants sold in the markets come from the wild, thus these places are inherently connected with the issue of sustainable collection of plant material from wild growing populations.
Rich traditions of the sale of plants in markets still persist in Europe, in spite of its high level of modernization. Probably the first regular studies of the ethnobotany of markets in the world, or at least in Europe, were performed by Polish scholars. In 1927 Muszyński [3] made a list of medicinal plants sold in the market of Vilnius (then Poland, nowadays Lithuania). Very soon after, in 1933, Jerzy Wojciech Szulczewski, a local biologist and ethnographer, issued a paper containing a detailed list of medicinal plants, and another about edible and medicinal fungi [4,5] (later reprinted in [6]) sold in the markets of Poznań, the largest city of western Poland. Just a few years later Szulczewski recorded 56 species of edible and medicinal fungi and 79 species of medicinal plants sold in these markets. His study was revisited in 2013 and a dramatic reduction in the sales of medicinal plants was observed [7].
Karousou et al. [8] studied medicinal herbs sold on 15 stalls scattered through markets in the three largest cities in Cyprus. A total of 57 taxa were recorded, of which 32 were cultivated and 14 wild. Similarly, Hanlidou et al. [9] studied medicinal plants in Thessaloniki, Greece. The majority (131) of the 172 recorded taxa were of local origin.
Łuczaj et al. [10] studied wild edible greens sold at 11 town markets in Dalmatia (on the southern coast of Croatia). According to the authors, the use of wild green vegetables (leaves, buds, stems) is very widespread in the Mediterranean. In total, 37 species were recorded.
Probably the longest list of plants sold in local markets was recorded by Ertug [11], in the Bodrum area of Turkey, who recorded 390 species on sale. Most of the recorded plants were wild edible plants, although plants used for fodder, medicine or crafts were also noted. Another study which recorded wild food plants in Turkey was carried out by Dogan et al. [12] who surveyed 18 markets in Izmir and found that 46 species of wild edible plants were sold. Nedelcheva and Dogan studied open-air markets on both sides of the Bulgarian-Turkish border. They found that predominantly medicinal plants are sold in Bulgarian markets, whereas in Turkish markets there are many more wild vegetables sold [13,14].
A number of scholars have researched plants sold in Asian markets. For instance Pemberton et al. [15] surveyed the three largest markets of wild edible and medicinal plants in Seoul, South Korea. Xu et al. [16] and X [17] looked at plants sold in Xishuangbanna, in the tropical part of Yunnan, China. Shirai & Rambo [18] presented the results of research on wild species sold on the main town square in Khon Kaen, in north -eastern Thailand. The diversity of wild species sold there is high; much higher in the rainy season than in the dry season. They found 60 wild species, of which 54 were plants, and 6 mushrooms. Konsam et al. [19] found a large diversity of wild vegetables sold in the markets of Manipur, India. The ethnobotany of open-air markets was also studied in Pakistan, Iraq, Iran and Kyrgyzstan [20][21][22][23].
One of the earliest ethnobotanical works on markets is by Bye [1] [26].
African markets are also dominated by medicinal plants (see e.g. [27][28][29][30]). Ouarghidi et al. [31] recorded medicinal plants sold in the three markets of Marrakech. They found that many of the species were falsifications and actually, false cheaper species are sold under different names. The fact that many important medicinal roots are not readily available in the markets of Marrakech suggests that these wild species may be in danger of extinction, and the scarcity and high demand for some species has led to their replacement by other taxa. The article lists species that are sold as replacements or forgeries. Similar results were obtained by Kool et al. [32] who found several rare and endangered species in the markets of southern Morocco.
In practically all of the above-mentioned studies from around the world the number of wild species sold is higher than the number of cultivated plants. Some authors raise the question to what extent such commerce endangers wild populations [33]. This is, for example, the case in Morocco concerning medicinal plants. The trade of ornamental plants, for example orchids, may also endanger local populations [34].
Another issue is the sale of mushrooms in open-air markets. It is widespread in many countries of the globe and often regulated [35][36][37], but the taxa which are sold are often not properly documented, due to the lack of voucher specimens. Recent advances in DNA barcoding techniques for fungi enable a more accurate identification of the species [38,39].
As previously mentioned, the issue of plant commodification is also connected with conservation statusplants are often protected because they are attractive, useful and prone to extinction. Although the first plant protected by law in Poland was the yew-tree (in 1420 by king Władysław Jagiełło - [40]), nowadays a large proportion of protected plants are those which have attractive flowers which make them prone to being picked or dug out for private gardens or for sale [41], for example Galanthus nivalis, Leucojum vernum, Orchidaceae, Lilium martagon etc. Such flowers formed the core of protected plants in the first post-World War II law on plant protection in Poland [42]. In Poland the overharvesting of wild medicinal plants used to be a problem, and rare medicinal plants constituted another large sector of protected plants. They were usually partially protected so that the authorities could license their gathering. Some of the species were not very rare at all (e.g. Asarum europaeum, Viburnum opulus, Frangula alnus) and their protection was cancelled in the most recent plant protection legislation [41], as the gathering of medicinal plants is much less widespread now. However, much earlier, at the end of the 19th century, many populations of highland alpine plants in the Tatras were decimated by pickers supplying medicinal "roots" to herbalists [43].
The aim of this study was to record wild plants and fungi sold in the markets of south-eastern Poland. Our hypotheses were:

Methods
The research was performed following the code of ethics of the American Anthropological Association [44] and the International Society of Ethnobiology Code of Ethics [45]. Oral prior informed consent was acquired. Four open-air markets from southeastern Poland were selected for the study (Fig. 1). This included probably the largest market in this part of Poland, in Rzeszów (190,000 inhabitants), the capital of Podkarpacie region. The other three markets were located in three (out of 21) county towns in the region -Jarosław (ca. 39,000 inhabitants), Leżajsk (14,000) and Przemyśl (63,000).
Open-air markets are located in most towns in Poland. In the past (e.g. before World War II) they were placed in a central market square (a typical feature of Polish medieval towns), but nowadays they were gradually relocated to less central locations. Once the centers of all commerce in towns, including selling animals, now they are a mixture of stalls with cheap clothes, tools, agricultural products, plants and mushrooms.
Data were collected in two ways. The first approach consisted of regular observations of markets. The markets of Rzeszów, Jarosław and Leżajsk ( Fig. 1) were visited every week on the days when most people come to buy products (J and L on Tuesday and R on Saturday) in the periods of September-October 2013, and from the end of February to the end of October 2014 and 2015. In 2015, the markets were visited only once every two weeks. Additionally the market in Przemyśl was visited seven times in 2015, from April to October, at monthly intervals. Altogether 13,488 photos were taken. Photographic documentation enabled the quick recording of plants used in public spaces and detailed identification of most taxa at least to genus level [46].
Photographs were taken on every visit in order to capture the diversity of cultivated and wild plants sold in the markets (Figs. 2, 3, 4, 5, 6, 7, 8 and 9). Lists of species sold were then made for each visit. Voucher specimens were collected if possible. Unfortunately, usually only parts of plants were available as vouchers and sometimes sellers refused to donate even parts of the plants they sold. The second part of the study consisted of interviews with the plant sellers. Altogether, 100 interviews were conducted (25 in each of the four markets). The interviews were carried out in the markets with the owners of plant stalls. The mean age of sellers was 55 There were 62 women and 38 men in the interviewed group. Around half of the sellers were farmers, the rest a variety of professionals, often retired. Profit and hobby were equally frequently cited motivations for selling plants in the market. The sellers had been selling plants for an average of 19 years (maximum selling time 50 years). As many as 62 % of sellers came every day, and the remainder 3-4 times a week.
The origins of the species (cultivation versus wild) were established based on interviews and the containers in which the species were sold. For example, species sold in small pots without any weeds and with wellestablished roots and one shoot were treated as cultivated, whereas species sold in plastic bags, dug out, with traces of natural vegetation (e.g. woodland mosses, other woodland or semi-natural grassland plants), were suspected to be collected from the wild (although they were sometimes dug out from gardens but then there were no traces of forest vegetation). In most cases the answers of the respondents were treated as trustworthy, with the exception of protected or believed-to-be-protected plants -sometimes sellers hesitated or gave very unclear answers when asked if the plant was dug out from the wild. In the case of strange behaviour from informants, we treated the plant as originating from the wild.   Most fungi specimens were successfully identified using the DNA barcoding technique [50,51]. Fungal DNA was extracted from a small part of each fruiting body using a Plant and Fungi DNA Purification Kit (Eurx), following standard protocol. The PCR cocktail was composed of 4 ml DNA extract, 0.5 ml each of the primers (ITS5 and ITS4 in 10 nmol concentration) and 5 ml Type-it Microsatellite PCR Kit (Qiagen). PCR was     [52], with the accession numbers listed in Table 1.

General information
We recorded 468 species of plants sold in the studied markets, including 117 species of plants which are native to Poland, or aliens (anthropophytes) established in the Podkarpacie region (Table 2). However the actual origins of native plants sold in the markets vary and included cultivated plants (84), plants which are both cultivated and collected from the wild (13 species), and species collected only from the wild (19 species). When only the plants which are at least partly collected from the wild are taken into account, most are sold for food, more rarely for medicinal and ornamental purposes (Fig. 10). When cultivated native plants are added, the ornamental purpose becomes dominant (Fig. 11), followed by food use and medicinal use. There are very few medicinal plants sold solely for this purpose (e.g. Tanacetum vulgae, Hypericum perforatum). Most medicinal plants are fruits used both for food and alcoholic drinks, and as medicine.
As many as 32 species of fungi are sold, all of them for culinary purposes (Table 3) Table 3).
The interviewed sellers mentioned 46 taxa of plants which they thought had been sold more often in the past, but they mentioned over twice as many (103 taxa, including non-native cultivated plants) as novelties, only recently sold. Out of the species which are sold less, the only wild/ native species was nettle Urtica dioica, mentioned by 2 informants. From the species which are seen as new or increasing, the informants mentioned a few native plants. Predominantly mentioned were ramsons Allium ursinum (5 informants), wormwood Artemisia absinthium (4), sorrel Rumex acetosa (3), Delphinium, blueberry Vaccinium spp. (2), sea buckthorn Hippophae rhamnoides and edelweiss Leontopodium alpinum (Table 4).

Protected plants
Relatively few protected plants (i.e. protected in the emic sense, according to respondents' knowledge) were listed as sold. Only 12 taxa of protected plants were mentioned by more than one informant. Allium ursinum, Convallaria maialis and Lycopodium were the most often cited. Most of the protected plants were cultivated in gardens (Table 1). This is especially the case with highland plants native only to the highest parts of the Carpathians (Leontopodium alpinum, Dryas octopetala etc.). The species which may be taken from the wild are Allium ursinum, club-mosses and orchids. (Un)fortunately club-mosses were not seen on sale     in the study period. The sellers of ramsons (Allium ursinum) claim they are cultivated but they have no certificate from local nature conservation authorities, and we suspect that the plants come from wild populations. Most of the protected plants are sold due to their ornamental value (apart from ramsons, whose use as vegetable has recently become fashionable). Poland is not the only country where plants with beautiful flowers are endangered. For example in Mexico the trade of ornamental orchids creates a biodiversity problem [34].
A very interesting issue is the difference between ordinary people's perceptions of protected plants and which plants are really protected. Our observations allow us to hypothesize that people are over-cautious about plant protection. There are some species which they think are protected but have never been, e.g. Anemone nemorosa. This wild species is common in the woods in Poland and many people think that it is protected by law because they learned at school about the protection of Anemone sylvestris and A. narcissiflora. Recently, in 2014, protection status was removed from many commoner species, which had previously been protected to avoid their being overharvested as medicinal plants (e.g. Frangula alnus, Viburnum opulus,    Asarum europaeum, Convallaria majalis), or because we now know that they are more common than previously thought (e.g. Equisetum telmateia).

Edible plants and fungi
Wild fruits make up an important sector of plants sold in the markets. From the interviews we infer that some species are re-appearing after years of neglect. The main reason for this re-emergence is the growing popularity of herbal medicine and fruit liqueurs. For example such plants as Rosa canina, Crataegus or Prunus spinosa are mainly sold for alcohol production, to a lesser extent also for herbal teas. This trend of the increasing availability of rarer economic plants in Polish market was already noticed by other authors [7,53]. Mushrooms constitute another important sector of the open-air markets. The number of recorded mushroom species is relatively high. It must be emphasized that the DNA bar-coding we used enabled confirmation of the identification of some surprising taxa sold in the markets (Lactarius quieticolor, Leccinum schistophilum, Leucoagaricus nympharum) never recorded as food in Poland before. Moreover, L. quieticolor and L. schistophilum are not listed in the checklist of Basidiomycetes found in Poland [54] and are new to the mycobiota of Poland. This highlights the importance of DNA barcoding in ethnomycological studies, illustrated well by the study of Dentinger and Suz [55] who found threw new species of porcini (Boletus) in a single packet of mushrooms sold in London and imported from China.
The list of fungi sold in the markets is similar to the taxa reported from a few ethnographic and ethnomycological studies in south-eastern Poland [56]. On the other hand some mushroom taxa widely collected in rural areas are very rarely sold, for example the Russula genus. Russulas can be confused with death cap Amanita phalloides, which is why they are not on the list of taxa permitted for sale in markets [37,57]. On the other hand Boletus luridiformis, which is not on the list of taxa allowed for commerce, is more frequently sold (and also traditionally collected here). Altogether 13 species of 32 sold in the markets are not on the list of species legally permitted for sale [57], and even one protected species [58] is sold (Table 3). In some cases very closely related taxa are listed in this legal document (Armillaria, Leccinum), in others the whole genus is not mentioned (even taking into account taxonomic changes and synonyms).
We may conclude that the choice of mushrooms in markets may be a good indicator of culturally salient edible mushrooms, though some lesser collected taxa may not be visible. A similar relationship was found between the traditionally collected wild vegetables in Dalmatia  [59], where there is high correlation between the two lists of taxa but some differences occur.

Comparison with other countries
It is difficult to compare Polish open-air markets with other European countries due to the scarcity of available data, which is restricted to south-eastern Europe. It seems that they have different features from the markets studied in Croatia, Bulgaria and the European part of Turkey. Turkish and Croatian markets sell a large number of wild green vegetables [10][11][12][13][14]. In Poland these are mainly restricted to Rumex species. Bulgarian markets, similarly to the Polish ones, have few wild vegetables, but contain many medicinal plants [13,14]. Such plants used to be sold in Polish markets in the first half of the 20th century [3][4][5][6]. Nowadays purely medicinal plants are rare in Polish markets but the category of culinary herbs is fashionable, however they are mainly nonnatives species. Such species as basil or oregano are relatively new to mainstream Polish cuisine and became fashionable a few years ago. People may be scared to sell purely medicinal plants due to regulations concerning the sales of medicinal material.

Conclusions
The open-air markets of southeastern Poland sell a considerable number of native plants but only a small proportion of them come from wild populations (mainly edible fruits). Most items are ornamental plants, or edible fruits and mushrooms. Very few medicinal plants and green vegetables are sold, which differentiates the markets from southern European ones. Such a pattern is probably the model for most central European markets, but no similar research has been carried out in neighbouring countries. Finding two species of fungi which are new to Poland highlights the importance of DNA barcoding in ethnomycological studies.