Unity in diversity—food plants and fungi of Sakartvelo (Republic of Georgia), Caucasus

Abstract

Background

The Republic of Georgia is part of the Caucasus biodiversity hotspot, and human agricultural plant use dates back at least 6000 years. Over the last years, lots of ethnobotanical research on the area has been published. In this paper, we analyze the use of food plants in the 80% of Georgia not occupied by Russian forces. We hypothesized that (1) given the long tradition of plant use, and the isolation under Soviet rule, plant use both based on home gardens and wild harvesting would be more pronounced in Georgia than in the wider region, (2) food plant use knowledge would be widely and equally spread in most of Georgia, (3) there would still be incidence of knowledge loss despite wide plant use, especially in climatically favored agricultural regions in Western and Eastern Georgia.

Methods

From 2013 to 2019, we interviewed over 380 participants in all regions of Georgia not occupied by Russian forces and recorded over 19,800 mentions of food plants. All interviews were carried out in the participants’ homes and gardens by native speakers of Georgian and its dialects (Imeretian, Rachian, Lechkhumian, Tush, Khevsurian, Psavian, Kakhetian), other Kartvelian languages (Megrelian, Svan) and minority languages (Ossetian, Ude, Azeri, Armenian, Greek).

Results

The regional division was based primarily on historic provinces of Georgia, which often coincides with the current administrative borders. The total number of taxa, mostly identified to species, including their varieties, was 527. Taxonomically, the difference between two food plant groups—garden versus wild—was strongly pronounced even at family level. The richness of plant families was 65 versus 97 families in garden versus wild plants, respectively, and the difference was highly significant. Other diversity indices also unequivocally pointed to considerably more diverse family composition of wild collected versus garden plants as the differences between all the tested diversity indices appeared to be highly significant.

The wide use of leaves for herb pies and lactofermented is of particular interest. Some of the ingredients are toxic in larger quantities, and the participants pointed out that careful preparation was needed. The authors explicitly decided to not give any recipes, given that many of the species are widespread, and compound composition—and with it possible toxic effects—might vary across the distribution range, so that a preparation method that sufficiently reduces toxicity in the Caucasus might not necessary be applicable in other areas.

Conclusions

Relationships among the regions in the case of wild food plants show a different and clearer pattern. Adjacent regions cluster together (Kvemo Zemo Racha, and Zemo Imereti; Samegrelo, Guria, Adjara, Lechkhumi and Kvemo and Zemo Svaneti; Meskheti, Javakheti, Kvemo Kartli; Mtianeti, Kakheti, Khevsureti, Tusheti. Like in the case of the garden food plants, species diversity of wild food plants mentioned varied strongly. Climate severity and traditions of the use of wild food plants might play role in this variation. Overall food plant knowledge is widely spread all-across Georgia, and broadly maintained.

Background

Georgia is situated between latitudes 41° and 44° N, and longitudes 40° and 47° E, with an area of 69,700 km², with 20% of the country currently occupied by Russian forces (Fig. 1). Georgia politically associates with European Union and takes part in all major programs of European development and cooperation. Georgia can be defined as a transcontinental country on the divide between Asia and Europe, with its larger part located south to this divide (i.e., in Asia) and smaller but strategically important parts (Khevi, Piriketi Khevsureti, etc.) located north of the continent divide (i.e., in Europe) [1].

Fig. 1.

Location and historical provinces of Georgia

The uplift of the Georgian Caucasus started in the late Oligocene and shares the same structural characteristics as the younger mountains of Europe. The Greater Caucasus mostly includes Cretaceous and Jurassic rocks, interspersed with Paleozoic and Precambrian formations in higher regions. Hard, crystalline, metamorphosed rocks like schist and gneisses, as well as pre-Jurassic granites are found in the western part, while softer, Early and Middle Jurassic clayey schist and sandstones in the eastern part. The foot of the Greater Caucasus are built of younger limestone, sandstones, and marls. The Lesser Caucasus in contrast is predominantly formed of Paleogene rocks interspersed with Jurassic and Cretaceous formations. The youngest geological structures of Georgia are represented by the vast volcanic plateaus in the southern part of country. These divisions lead to an extremely complex terrain with pronounced climatic gradients: (1) the mountains of the greater Caucasus with peaks over 5000 m (Shkara, Babis Mta, Chanchakhi, etc.); (2) the inter-mountain plains between the Greater and Lesser Caucasus mountains; (3) the mountains of the Lesser Caucasus with peaks rarely exceeding 3000 m (Mepistskaro, Kheva, Shavi Klde, Kanis Mta, Arsiani); (4) the Volcanic plateau of the Southern Georgia with elevations from 1300 to 2200 m [2–4].

Georgia’s climate is influenced by its location in the warm temperate zone stretching from the Black to the Caspian Seas, and the complexity of its terrain. Georgia has a coastline of 330 km with warm climate, the mean temperature reaching 4–7 °C in January and 22–23 °C in July, and high precipitation (1500–2000 mm annually). The warm oceanic-subtropical climate can be found only at lower elevations (less than 650 m); in more elevated terrains and to the north and east the climate becomes moderately warm. The Greater Caucasus bars cold air from the north, while warm and moist air from the Black Sea spreads easily into the coastal lowlands until reaching the Likhi range, which partly impedes further westward movement of the warm and moist air. In central Georgia, precipitation in mountains can be twice that in the plains. In the mountains, weather conditions change to cool and wet quite steeply with increasing elevation and above 2100 m the environment becomes sub-alpine and alpine, with permanent snow and ice above 3600 m [2–4].

Plant use history

The Caucasus is regarded as global biodiversity hotspot [5–8]. Botanical has a long history, and the vegetation composition as well as flora are well-known [2, 3].

The territory of modern-day Georgia (Fig. 1) has been inhabited since the early Stone Age, and agriculture was already well-developed during the early Neolithic [9], although human occupation started already in the Early Pleistocene, with the 1.7-Myr-old hominid fossils of Dmanisi in Southern Georgia being the earliest known hominid-site outside of Africa [10–12]. The history of plant and animal use has been documented since the Upper Paleolithic through fossils found in Dzudzuana Cave, dated to ~ 36–34 Ka BP, including wool (Capra caucasica), and dyed fibers of wild flax (Linum usitatissimum) [13]. Archeological findings from the Neolithic and Early Bronze periods dating back to the 6th–2nd millennium BC are rich with plant fossils and seeds of both wild species and local landraces [14]. The earliest seeds of Vitis vinifera (grapevine) were excavated in southern Georgia and date to about 8000 years BP [15]. Medicinal species like Alchemilla millefolium, Artemisia annua, A. absinthium, Centaurea jacea and Urtica dioica found in the archeological record are still used in the modern pharmacopoeia [16].

Due to its ancient roots agriculture in Georgia is characterized by a great diversity of landraces, and endemic species of crops, already documented in Soviet times [17–22]. However, starting with the implementation of Stalinist agricultural reforms in the 1950s, a rapid loss of local cultivars occurred [23–26]. This process accelerated during post-independence, and knowledge loss has been shown to even extent to aggravate wolf-human conflicts [27]. However, a wide variety of local cultivars can still be found in case of Vitis vinifera (Vitaceae) shows its highest genetic diversity in Georgia, with over 600 varieties known, and several dozen used commercially [9, 15, 28–31]. In contrast, essentially none of the 144 varieties, and 150 forms of wheat (Triticum) registered in Georgia in the 1940s [21, 22] are sown in modern Georgian commercial agriculture [25], although traditional varieties are still reported from nearby Turkey [32]. The situation is similar in case of Hordeum vulgare (Poaceae) which originally was important in beer production, for religious rituals and traditional medicine [9, 33] and Secale cereale (Poaceae) [34].

In contrast to the loss of cereals, legumes like peas (Pisum sativum), lentils (Lens cornicularis), chickpeas (Cicer arietinum), fava beans (Vicia faba), and vegetables like garden lettuce (Lactuca sativa), beans (Phaseolus vulgaris), basil (Ocimum basilicum), peppermint (Mentha x piperita), onions (Allium cepa), sugar beets (Beta vulgaris), spinach (Spinacia oleracea), carrots (Daucus carota), radishes (Raphanus sativus), turnips (Brassica rapa var. rapa), welsh onion (Allium fistulosum), amaranth (Amaranthus viridis), goosefoot (Chenopodium album), leeks (Allium ampeloprasum) and garlic (Allium sativum) are still common in home gardens. Herbs like parsley (Petroselinum crispum), coriander (Coriandrum sativum), tarragon (Artemisia dracunculus), savory (Satureja hortensis), garden cress (Lepidium sativum), dill (Anethum graveolens), fennel (Foeniculum vulgare), celery (Apium dulce), and Allium fistulosum (Chinese onion are widely cultivated and popular ingredients of local cuisine [1]. The maintenance of such diversity is of high importance as source material for global crop production [35, 36]. Many species are widely sold as medicines [37].

Over the last years, ethnobotanical research in Georgia has received a large boost, and a wide variety of studies on all aspects of plant use have been published [38–52]. Few of these however focused entirely of food plants [38, 52], many of which are still cultivated in small home-gardens. Home-gardens are often cited as important reservoirs for crop germplasm [53–58] and are mostly sources of food [59, 60]. In wider Eurasia, home gardens have been shown to be an important repository of plant diversity are often part of complex seed exchange networks [61–64].

Given the trajectory of ethnobotanical studies in Georgia, a meta-analysis of the data food plant uses was long overdue. In this publication, we hypothesized that (1) given the long tradition of plant use, and the isolation under Soviet rule, plant use both based on home gardens and wild harvesting would be more pronounced in Georgia than in the wider region, (2) food plant use knowledge would be widely and equally spread in most of Georgia, (3) there would still be incidence of knowledge loss despite wide plant use, especially in climatically favored agricultural regions in Western and Eastern Georgia.

Materials and methods

Ethnobotanical interviews

From 2013 to 2019, we interviewed over 380 participants in all regions of Georgia not occupied by Russian forces on their general plant use, recording over 32,000 individual uses. The analyses of all uses have been published in a variety of papers [41–50]. However, of all uses over 19,800 mentions were of food plants, which is why we regarded it as prudent to present a separate analysis of these. Interviews using semi-structured questionnaires were conducted after obtaining the oral prior informed consent of the participants, which were selected by snowball sampling, trying to reach gender balance and representing different age groups. Most participants were however over 50 years old, as interviews targeted remote villages where only very few younger people remain. All interviews were carried out in the participants’ homes and gardens by native speakers of Georgian and its dialects (Imeretian, Rachian, Lechkhumian, Tush, Khevsurian, Psavian, Kakhetian), other Kartvelian languages (Megrelian, Svan) and minority languages (Ossetian, Ude, Azeri, Armenian, Greek). The languages in which a plant was mentioned are indicated in Table 1. Interviews were subsequently translated into English. Plants grown in home gardens were used as prompts, while wild-collected species were free listed. We classified species as "garden" when they were grown/collected in cultivated areas, and as "forest/wild-collected" when growing and harvested in the wild. We maintained the distinction of "forest" and "garden" because it was used in our previous publications from the region [50], to maintain consistency. In contrast to many other countries Georgia benefits from a complete flora [65–69] and a broad inventory of vernacular names in all languages [68]. Species were identified directly in the field, using this literature, and vouchers collected and deposited in the National Herbarium of Georgia (TBI). The nomenclature of all species follows www.tropicos.org, under APGIII [70]. Collection permits were provided through the Institute of Botany, Ilia State University, Tbilisi.

Table 1.

All Food plant and fungal species encountered in Georgia

Data analysis

Data were tabulated using excel sheets and a combined matrix was constructed with plant entries in rows and plant data in columns including date, place, participant’s age and gender, interviewer, plant identity (Latin, Georgian vernacular, local names), the use category, which parts were used, and the source (garden or forest). We compared species diversity among groups of species (forest versus garden, various provinces) using sample-based rarefaction as well as widely used diversity indices: Dominance (D), Shannon (H), Evenness (e^H/S), Simpson index, (1 − D), Equitability (J), Fisher alpha, Berger–Parker (BP), given that no single index may sufficiently show the importance of certain species. Similarity of species composition among groups of plants were analyzed using non-metric multidimensional scaling (nMDS). All these analyses were performed using software PAST4.02 [71].

Test if the usage of plants based on family and genus, plant system used, and general and specific plant parts differ between regions and different altitudinal ranges. I predict that these components will be different, since there will be a different plant composition among regions and along an altitudinal gradient, and that different human communities have their own ethnobotany knowledge, even though they are from the same country.

We compared the usage of plants based on their (i) family and (ii) genus, (iii) system (root, shoot, or both), and (iv) general (vegetative, reproductive, or both) and (v) more specific (bark, branches, buds, bulb, cones, flowers, fruit, latex, leaves, resin, roots, seeds, shoots, silk, stem, timber, tuber, whole plant) parts used between regions and altitudinal ranges. We also compared (vi) for what purpose plants are used between regions and altitudinal ranges. We removed from our analyses any data that was not possible to make any further identification, such as plants identification above family, and uncertain plant parts. We also removed fungi from our analyses, and samples in which we had no more details about the purpose of usage of plants, i.e., in cases where plants were used as human food, but we did not know exactly for which kind of food. We considered regions and five altitudinal ranges (0–500 m, 501–1000 m, 1001–1500 m, 1501–2000 m, 2001–2500 m) as factors within our ordinations. We conducted non-metric multidimensional scaling (NMDS) followed by a permutational multivariate analysis of variance (PERMANOVA) with Euclidean distance and 999 permutations using the “RVAideMemoire” package [72].

Results

The total number of taxa, mostly identified to species, was 527 (Tables 1 and 2, Appendix Tables 5, 6). Ninety-five species of fungi were consumed. Trees contributed 71 species (13.47%), Shrubs—43 (8.1%), Herbs—333 (60.32%), Climbers -5 (0.09%), and Fungi—95 (18.02%). Of all species 388 were wild, i.e., not cultivated, although some of them occurred on ruderal places and as weeds in gardens. In case of 20 vascular plants and 45 fungal species, the collected material did not allow a certain identification, and these species are thus indicated as "indet." in Table 1. Taxonomically, the difference between two food plant groups—garden versus wild ("forest")—was strongly pronounced even at family level. Only one plant species (Piper nigrum with four mentions) was bought in markets. Over 62% of the mentions (12,255) referred to cultivated plants, 7352 (37%) to wild collections, and some plants were found both collected in the wild and in gardens; however, this was a very small percentage (189 mentions, less than 1%). The great majority of mentions (> 99%) were either from families found either in gardens (62%) or in the wild (37%). Over 41% of all mentions referred to the use of fruits, 21% to leaves, about 7% to seeds, and 5% to fruiting bodies, leaves/stems and stems. Whole plants were only used very infrequently. Of all the families, Rosaceae, Apiaceae, Lamiaceae, Amaryllidaceae and Solanaceae showed the highest importance. At a generic level, Allium, Pyrus, Malus and Brassica received the highest number of use report. Only 30 species (6% of the total) represented 46% of all use mentions, but only Malus orientalis (3.5%), Pyrus communis (3.2%), and Vitis vinifera (2.7%) had over 2% of mentions, and Chenopodium album and Urtica dioica were the only not cultivated plants reaching over 1% of mentions. In most regions at all altitudinal ranges, the aboveground parts were mist frequently used (Fig. 2),

Table 2.

Regions of our fieldwork and number of food plant mentions recorded

Region	Number of mentions
Guria	2125
Khevsureti	2012
Zemo Svaneti	1942
Adjara	1866
Tori	1750
Tusheti	1633
Kvemo Svaneti	1406
Kakheti	1085
Lechkhumi	1017
Samegrelo	853
Meskheti	776
Kvemo Racha	708
Javakheti	699
Kvemo Kartli	678
Zemo Imereti	631
Mtianeti	342
Zemo Racha	277

Table 5.

Species of identified food plants and fungi and the number of their mentions recorded

Plant / Fungal family	Plant / Fungal species	Mentions
Actinidiaceae	Actinidia callosa Lindl	28
Adoxaceae	Sambucus ebulus L	83
Adoxaceae	Sambucus nigra L	9
Adoxaceae	Viburnum lantana L	21
Adoxaceae	Viburnum opulus L	21
Agaricaceae	Agaricus arvensis Schaeff	165
Agaricaceae	Agaricus campestris L	4
Agaricaceae	Agaricus tabularis Peck	1
Agaricaceae	Bovista sp.	12
Agaricaceae	Bovista sp. / Lycoperdon sp.	4
Agaricaceae	Clavatia gigantea (Batsch) Rostk	14
Agaricaceae	Coprinus comatus (O.F. Müll.) Pers	2
Agaricaceae	Lycoperdon perlatum Pers. / Lycoperdon pyriforme Schaeff	2
Amanitaceae	Amanita caesarea (Scop.) Pers	15
Amanitaceae	Amanita muscaria (L.) Lam	1
Amaranthaceae	Amaranthus palmeri S. Watson	16
Amaranthaceae	Amaranthus paniculatus L	24
Amaranthaceae	Amaranthus retroflexus L	132
Amaranthaceae	Amaranthus speciosus L	1
Amaranthaceae	Amaranthus spinosus L	3
Amaranthaceae	Atriplex hortensis L	35
Amaranthaceae	Beta vulgaris L	311
Amaranthaceae	Beta vulgaris L. ssp. cicla (L.) Moq	36
Amaranthaceae	Beta vulgaris L. ssp. esculenta (Salisb.) Gürke var. altissima Rössig. = Beta vulgaris saccharifera Alef	3
Amaranthaceae	Chenopodium album L	203
Amaranthaceae	Chenopodium bonus-henricus L	1
Amaranthaceae	Chenopodium foliosum (Moench) Asch	35
Amaranthaceae	Chenopodium sp.	1
Amaranthaceae	Spinacia oleracea L	44
Amaryllidaceae	Allium ampeloprasum L	3
Amaryllidaceae	Allium ascalonicum L	7
Amaryllidaceae	Allium atroviolaceum Boiss	10
Amaryllidaceae	Allium cepa L	309
Amaryllidaceae	Allium fistulosum L	97
Amaryllidaceae	Allium kunthianum Vved	2
Amaryllidaceae	Allium ponticum Miscz	5
Amaryllidaceae	Allium porrum L	56
Amaryllidaceae	Allium rotundum L	20
Amaryllidaceae	Allium sativum L	340
Amaryllidaceae	Allium sp.	3
Amaryllidaceae	Allium ursinum L	54
Amaryllidaceae	Allium victorialis L	231
Amaryllidaceae	Galanthus sp.	10
Amaryllidaceae	Galanthus woronowii Losinsk	3
Amaryllidaceae	Narcissus sp.	5
Annonaceae	Annona cherimola Mill	1
Apiaceae	Aethusa cynapium L	1
Apiaceae	Agasyllis latifolia (Bieb.) Boiss	91
Apiaceae	Anethum graveolens L	301
Apiaceae	Angelica tatianae Bordz	2
Apiaceae	Anthriscus cerefolium (L.) Hoffm	4
Apiaceae	Anthriscus nemorosus (M. Bieb.) Spreng	16
Apiaceae	Anthriscus sylvestris L	15
Apiaceae	Apium graveolens L	128
Apiaceae	Carum carvi L	60
Apiaceae	Chaerophyllum aureum L	16
Apiaceae	Chaerophyllum bulbosum L	10
Apiaceae	Chaerophyllum caucasicum (Fisch.) B. Schischk	95
Apiaceae	Conium maculatum L	10
Apiaceae	Coriandrum sativum L	348
Apiaceae	Daucus carota L. ssp. sativus	251
Apiaceae	Falcaria sioides Asch	1
Apiaceae	Falcaria vulgaris Bernh	25
Apiaceae	Foeniculum vulgare Mill	79
Apiaceae	Heracleum asperum M. Bieb	30
Apiaceae	Heracleum leskovii Grossh	5
Apiaceae	Heracleum sect. villosum	2
Apiaceae	Heracleum sosnowskyi Manden	59
Apiaceae	Heracleum sp.	36
Apiaceae	Heracleum wilhelmsii Fisch. & Ave-Lall	30
Apiaceae	Hippomarathrum crispum (Pers.) Boiss	4
Apiaceae	Hippomarathrum microcarpum Petrov	1
Apiaceae	Levisticum officinale W.D.J. Koch	2
Apiaceae	Libanotis transcaucasica Schischk	15
Apiaceae	Ligusticum alatum Spreng	4
Apiaceae	Petroselinum crispum (Mill.) Fuss	268
Apiaceae	Xanthogalum purpurascens Avé-Lall	3
Araceae	Arum albispatum Stev. ex Ledeb	2
Araceae	Arum orientale M. Bieb	7
Araceae	Arum sp.	20
Araliaceae	Aralia spinosa L	1
Asparagaceae	Asparagus officinalis L	30
Asparagaceae	Asparagus sp.	4
Asparagaceae	Muscari sosnowskyi Schchian	2
Asparagaceae	Ornithogalum woronowii Kasch	2
Asparagaceae	Polygonatum glaberrimum C. Koch	13
Asparagaceae	Ruscus colchicus Yeo	1
Asparagaceae	Ruscus hypophyllum L	2
Asparagaceae	Scilla siberica Andrews	6
Asparagaceae	Scilla sp.	6
Asteraceae	Achillea grandiflora M. Bieb	1
Asteraceae	Achillea millefolium L	5
Asteraceae	Arctium lappa L	32
Asteraceae	Artemisia absinthium L	8
Asteraceae	Artemisia dracunculus L	125
Asteraceae	Artemisia vulgaris L	3
Asteraceae	Bidens tripartida L	4
Asteraceae	Cichorium intybus L	11
Asteraceae	Cirsium incanum (S.G. Gmel.) Fisch. ex M. Bieb	13
Asteraceae	Cirsium sp.	5
Asteraceae	Cirsium vulgare L	3
Asteraceae	Crepis sp.	3
Asteraceae	Cynara cardunculus L	6
Asteraceae	Echinops sp.	2
Asteraceae	Eruca sativa Mill	12
Asteraceae	Helianthus annuus L	17
Asteraceae	Helianthus tuberosus L	17
Asteraceae	Lactuca sativa L	165
Asteraceae	Lactuca sativa L. "greek"	1
Asteraceae	Lactuca serriola L	17
Asteraceae	Lapsana communis L	9
Asteraceae	Lapsana grandiflora M. Bieb	2
Asteraceae	Matricaria chamomilla L	5
Asteraceae	Petasites albus (L.) Gaertn	14
Asteraceae	Petasites hybridus (L.) G. Gaert, B. Mey. & Scherb	51
Asteraceae	Serratula quinquefolia Bieb. ex Willd	20
Asteraceae	Solidago canadensis L	4
Asteraceae	Sonchus asper (L.) Hill	7
Asteraceae	Stevia sp.	2
Asteraceae	Tagetes patula L	114
Asteraceae	Taraxacum confusum Schischk	2
Asteraceae	Taraxacum officinale Wigg	41
Asteraceae	Tragopogon sp.	19
Asteraceae	Tussilago farfara L	1
Asteraceae	Xanthium strumarium L	3
Auriculariaceae	Auricularia auricula-judae (Bull.) Quél	10
Bankeraceae	Hydnum repandum Fr	2
Bankeraceae	Sarcodon imbricatus (L.) P. Karts	8
Begoniaceae	Begonia rex Putz	10
Berberidaceae	Berberis vulgaris L	54
Betulaceae	Alnus barbata C.A. Mey	1
Betulaceae	Betula litwinowii Doluch	3
Betulaceae	Betula sp.	2
Betulaceae	Corylus avellana L. / C. pontica K. Koch	200
Betulaceae	Corylus iberica L	4
Boletaceae	Boletus edulis Bull	16
Boletaceae	Neoboletus erythropus (Pers.) C. Hahn	2
Boletaceae	Leccinum scabrum (Bull.) Gray	3
Boraginaceae	Myosotis sp.	2
Boraginaceae	Symphytum grandiflorum DC	14
Boraginaceae	Trachystemon orientalis (L.) G. Don	6
Brassicaceae	Armoracia rusticana (G. Gaertn.) B. Mey. & Scherb	33
Brassicaceae	Brassica campestris L	1
Brassicaceae	Brassica campestris L. ssp. oleifera DC	9
Brassicaceae	Brassica juncea (L.) Czern	3
Brassicaceae	Brassica montana Pourr	36
Brassicaceae	Brassica oleracea L	361
Brassicaceae	Brassica oleracea L. red	9
Brassicaceae	Brassica oleracea L. var. botrytis cauliflower	25
Brassicaceae	Brassica oleracea L. var. gemmifera Brussles Sprouts	1
Brassicaceae	Brassica oleracea L. var. gongylodes	47
Brassicaceae	Brassica oleracea L. var. italica	21
Brassicaceae	Brassica rapa L. subsp. rapifera Metzger	67
Brassicaceae	Brassica rapa var. rapa L	45
Brassicaceae	Bunias orientalis L	27
Brassicaceae	Capsella bursa-pastoris L	26
Brassicaceae	Cardamine hirsuta L	10
Brassicaceae	Cheiranthus cheiri L	1
Brassicaceae	Lepidium sativum L	52
Brassicaceae	Raphanus raphanistrum subsp. sativus (L.) Domin	17
Brassicaceae	Raphanus sativus L. var. major	179
Brassicaceae	Raphinastrum rugosum L. All	13
Brassicaceae	Sinapis arvensis L	15
Campanulaceae	Campanula alliariifolia Wild	2
Campanulaceae	Campanula biebersteiniana Roem. & Schult	1
Campanulaceae	Campanula glomerata L	7
Campanulaceae	Campanula lactiflora M. Bieb	70
Campanulaceae	Campanula latifolia L	11
Campanulaceae	Campanula rapunculoides L	20
Cannabaceae	Cannabis sativa L	30
Cannabaceae	Humulus lupulus L	22
Cantharellaceae	Cantharellus cibarius Fr	36
Caprifoliaceae	Lonicera caucasica Pall	3
Caryophyllaceae	Melandrium balansae Boiss	5
Caryophyllaceae	Melandrium boissieri Schischk	9
Caryophyllaceae	Melandrium sp.	5
Caryophyllaceae	Oberna wallichiana (Klotzsch) Ikonn	3
Caryophyllaceae	Silene lacera Steven	15
Caryophyllaceae	Silene sibirica (L.) Pers	2
Caryophyllaceae	Silene wallachiana Klotzsch	9
Caryophyllaceae	Stellaria media (L.) Vill	9
Clavariadelphaceae	Clavariadelphus pistillaris (L.) Donk	5
Convolvulaceae	Convolvulus arvensis L	17
Cornaceae	Swida australis (C.A. Mey.) Pojark ex Grossh	5
Cortinariaceae	Cortinarius violaceus (L.) Fr. Gray	1
Crassulaceae	Sedum caucasicum Boriss	8
Crassulaceae	Sedum oppositifolium Sims	5
Crassulaceae	Sedum stoloniferum Gmel	5
Crassulaceae	Sempervivum caucasicum Rupr. ex Boiss	14
Cucurbitaceae	Bryonia dioica Jacq	3
Cucurbitaceae	Citrullus lanatus (Thunb.) Matsum. & Nakai	16
Cucurbitaceae	Cucumis melo L	4
Cucurbitaceae	Cucumis sativus L	363
Cucurbitaceae	Cucurbita maxima L	14
Cucurbitaceae	Cucurbita pepo L	201
Cucurbitaceae	Cucurbita pepo L. var. giromontia	39
Cucurbitaceae	Cucurbita pepo L. var. patisson	9
Cucurbitaceae	Cucurbita sp.	14
Cucurbitaceae	Lagenaria siceraria (Molina) Standl	2
Cupressaceae	Junperus sabina L	2
Dipsacaceae	Cephalaria gigantea (Ledeb.) Bobrov	1
Dryopteridaceae	Dryopteris filix-mas (L.) Schott	35
Ebenaceae	Diospyros lotus L	54
Ebenaceae	Diospyros sp.	4
Ebenaceae	Diospyros virginiana L	5
Elaeagnaceae	Elaeagnus sp.	3
Elaeagnaceae	Hippophaë rhamnoides L	3
Elaeagnaceae	Shepherdia argentea Nutt	1
Elaeagnaceae	Shepherdia sp.	3
Ericaceae	Empetrum hermaphroditum Hagerup	21
Ericaceae	Oxycoccus quadripetalus Gilib	1
Ericaceae	Vaccinium arctostaphylos L	190
Ericaceae	Vaccinium myrtillus L	209
Ericaceae	Vaccinium sp.	4
Ericaceae	Vaccinium uliginosum L	2
Ericaceae	Vaccinium vitis-idaea L	49
Euphorbiaceae	Aleurites moluccanua (L.) Willd	1
Fabaceae	Astragalus caucasisus Pall	1
fabaceae	Cicer arietinum L	25
Fabaceae	Coronilla varia L	5
Fabaceae	Galega orientalis Lam	9
Fabaceae	Glycine max (L.) Merr	35
Fabaceae	Glycyrrhiza glabra L	1
Fabaceae	Lathyrus roseus Steven	42
Fabaceae	Lathyrus tuberosus L	3
Fabaceae	Lens cornicularis L	16
Fabaceae	Phaseolus sativus L	270
Fabaceae	Phaseolus vulgaris L	86
Fabaceae	Pisum sativum L	66
Fabaceae	Robinia pseudoacacia L	45
Fabaceae	Trifolium sp.	5
Fabaceae	Trigonella caerulea (L.) Ser	173
Fabaceae	Vicia faba L	54
Fabaceae	Vicia sativa L	1
Fabaceae	Vigna angularis (Willd.) Ohwi & H. Ohashi	1
Fagaceae	Castanea sativa Mill	79
Fagaceae	Fagus orientalis Lipsky	53
Fagaceae	Quercus iberica M. Bieb	9
Fistulinaceae	Fistulina hepatica (Schaeff.) With	6
Fungi	Unidentified fungus	227
Gentianaceae	Swertia iberica Fisch & C.A. Mey	1
Geraniaceae	Erodium cicutarium (L.) L'Hér. ex Aiton	4
Geraniaceae	Geranium robertianum L	3
Geraniaceae	Geranium sp.	6
Grossulariaceae	Grossularia reclinata (L.) Mill	27
Grossulariaceae	Ribes biebersteinii Berl. ex DC	59
Grossulariaceae	Ribes grossularia L	22
Grossulariaceae	Ribes nigrum L	73
Grossulariaceae	Ribes orientale Desf	4
Grossulariaceae	Ribes rubrum L	103
Grossulariaceae	Ribes sp.	24
Grossulariaceae	Ribes uva-crispa L	13
Guttiferae	Hypericum perforatum L	22
Hericiaceae	Hericium erinaceus (Bull.) Pers	1
Iridaceae	Crocus sativus L	9
Juglandaceae	Juglans mandshurica Maxim	7
Juglandaceae	Juglans regia L	235
Juglandaceae	Pterocarya pterocarpa (Michx.) Kunth ex Iljinsk	7
Lamiaceae	Lamium album L	32
Lamiaceae	Lamium purpureum L	6
Lamiaceae	Leonotis leonurus (L.) R. Br	1
Lamiaceae	Mentha aquatica L	3
Lamiaceae	Mentha longifolia (L.) L	158
Lamiaceae	Mentha pulegium L	81
Lamiaceae	Mentha sp.	8
Lamiaceae	Mentha x piperita L	143
Lamiaceae	Nepeta mussinii Spreng	2
Lamiaceae	Ocimum basilicum L	198
Lamiaceae	Ocimum basilicum var. purpurascens Benth	8
Lamiaceae	Origanum vulgare L	50
Lamiaceae	Salvia verticillata L	3
Lamiaceae	Satureja hortensis L	92
Lamiaceae	Satureja laxiflora K. Koch	7
Lamiaceae	Satureja spicigera Boiss	31
Lamiaceae	Thymus caucasicus Willd. ex Benth	30
Lamiaceae	Thymus colinus Bieb	21
Lamiaceae	Thymus sp.	29
lamiaceae	Thymus transcaucasicus Ronninger	17
Lamiaceae	Ziziphora pushkinii Adams	18
Lamiaceae	Ziziphora serpyllacea M. Bieb	16
Lauraceae	Laurus nobilis L	25
Lauraceae	Persea americana Mill	2
Lepiotaceae	Macrolepiota procera (Scop.) Springer	51
Liliaceae	Fritillaria lutea Mill	11
Liliaceae	Gagea sp.	3
Liliaceae	Lilium sp.	1
Liliaceae	Lilium szovitsianum Fisch. & Avé-Lall	11
Liliaceae	Ornithogalum woronowii Kasch	6
Linaceae	Linum usitatissimum L	7
Lythraceae	Punica granatum L	32
Malvaceae	Alcea rosea L	1
Malvaceae	Althaea spp.	11
Malvaceae	Malva neglecta L	38
Malvaceae	Malva sylvestris L	10
Malvaceae	Malva sylvestris L. / M. neglecta L	59
Malvaceae	Tilia begonifolia Stev	2
Malvaceae	Tilia caucasica Rupr	49
Marasmiaceae	Marasmius oreades (Bolton) Fr	12
Melanthiaceae	Veratrum lobelianum Bernh	5
Moraceae	Ficus carica L	142
Moraceae	Morus alba L	99
Moraceae	Morus nigra L	7
Morchellaceae	Morchella conica Pers	1
Morchellaceae	Morchella esculenta (L.) Pers	12
Musaceae	Musa x paradisiaca L	3
Myrtaceae	Acca sellowiana (O. Berg.) Burret	11
Oleaceae	Fraxinus excelsior L	5
Oleaceae	Ligustrum vulgare L	2
Onagraceae	Chamenaerion angustifolium (L.) Holub	1
Onocleaceae	Mattheuccia struthiopteris (L.) Todd	35
Orobanchaceae	Pedicularis sp.	5
Oxalidaceae	Averrhoa carambola L	1
Oxalidaceae	Oxalis acetosela L	1
Oxalidaceae	Oxalis corniculata L	1
Papaveraceae	Papaver somniferum L	32
Physalacriaceae	Armillariella mellea (Vahl) P. Kumm	93
Phytolaccaceae	Phytolacca americana L	12
Pinaceae	Abies nordmanniana (Steven) Spach	7
Pinaceae	Cedrus sp.	3
Pinaceae	Picea orientalis (L.) Peterm	17
Pinaceae	Pinus kochiana Klotzsch ex K. Koch	10
Pinaceae	Pinus sosnowskyi Nakai	8
Piperaceae	Piper nigrum L	4
Plantaginaceae	Plantago major L	2
Plantaginaceae	Valeriana officinalis L	1
Pleurotaceae	Pleurotus cornicopiae (Paulet) Rolland	4
Pleurotaceae	Pleurotus ostreatus (Jacq. ex Fr.) P. Kumm	90
Pluteaceae	Pluteus cervinis (Schaeffer ex Fr). P. Kumm	28
Poaceae	Avena sativa L	42
Poaceae	Bambusa sp.	4
Poaceae	Hordeum vulgare L	97
Poaceae	Hordeum vulgare L. ssp. vulgare L. var. coelestre L	5
Poaceae	Panicum crus-calli L	2
Poaceae	Panicum milanjianum Rendle	38
Poaceae	Secale cereale L	65
Poaceae	Setaria italica (L.) P. Beauv	16
Poaceae	Sorghum bicolor (L.) Moench	2
Poaceae	Triticum aestivum L	144
Poaceae	Triticum carthlicum Nevski	4
Poaceae	Triticum dicoccum Schrank	2
Poaceae	Triticum sp.	2
Poaceae	Zea mays L	195
Polygonaceae	Fagopyrum tataricum (L.) Gaertn	9
Polygonaceae	Polygonum alpinum All	57
Polygonaceae	Polygonum aviculare L	9
Polygonaceae	Polygonum carneum C. Koch	74
Polygonaceae	Polygonum panjutini Kharkev	5
Polygonaceae	Polygonum sp.	6
Polygonaceae	Rheum rhabarbarum L	3
Polygonaceae	Rumex acetosa L	77
Polygonaceae	Rumex acetosella L	19
Polygonaceae	Rumex alpinus L	84
Polygonaceae	Rumex crispus L	44
Polygonaceae	Rumex scutatus L	6
Polygonaceae	Rumex sp.	20
Polygonaceae	Rumex tuberosus L	1
Polypodiaceae	Polypodium vulgare L	10
Polyporaceae	Polyporus squamosus (Huds.) Fr	9
Portulacaceae	Portulaca oleracea L	85
Primulaceae	Cyclamen vernum Sweet	5
Primulaceae	Primula luteola Rupr	1
Primulaceae	Primula macrocalyx Bunge	24
Primulaceae	Primula sp.	4
Primulaceae	Primula vulgaris subsp. rubra (Sm.) Arcang	3
Primulaceae	Primula woronowii Losinsk	18
Psathyrellaceae	Coprinopsis atramentaria (Bull.) Redhead, Vilgalys & Moncalvo	24
Ramariaceae	Ramaria flava (Schaeff.) Quél	18
Ranunculaceae	Adonis aestivalis L	2
Ranunculaceae	Clematis vitalba L	11
Ranunculaceae	Ranunculus repens L	2
Rhamnaceae	Rhamnus imeretina Booth, Petz. & Kirchn	1
Rhamnaceae	Ziziphus jujuba Mill	2
Rhododendraceae	Rhododendron caucasicum Pall	79
Rhododendraceae	Rhododendron luteum Sweet	15
Rhododendraceae	Rhododendron ponticum L	27
Rosaceae	Armeniaca vulgaris Lam	2
Rosaceae	Aruncus vulgaris Raf	31
Rosaceae	Cornus mas L	135
Rosaceae	Cotoneaster multiflorus Bunge	4
Rosaceae	Crataegus curvisepala Lindm	34
Rosaceae	Crataegus pentagyna Waldst	48
Rosaceae	Crataegus sp.	13
Rosaceae	Cydonia oblonga L	80
Rosaceae	Duchesnea indica (Andrews) Teschem	6
Rosaceae	Eriobotrya japonica (Thunb.) Lindl	27
Rosaceae	Fragaria vesca L	74
Rosaceae	Fragaria vesca L. "Alibaba"	1
Rosaceae	Fragaria virginiana Mill	12
Rosaceae	Fragaria x ananassana Duchesne ex Rozier	35
Rosaceae	Malus orientalis Uglizk	685
Rosaceae	Malus pumila Mill. var. paradisiaca C.K. Schneid	3
Rosaceae	Mespilus germanica L	81
Rosaceae	Padus racemosa (Lam.) Gilib	27
Rosaceae	Prunus amygdalus Batsch	1
Rosaceae	Prunus armeniaca L	30
Rosaceae	Prunus avium (L.) L	187
Rosaceae	Prunus cerasus L	78
Rosaceae	Prunus divaricata Ledeb	282
Rosaceae	Prunus insititia L	62
Rosaceae	Prunus laurocerasus L	63
Rosaceae	Prunus padus L	2
Rosaceae	Prunus persica (L.) Batsch	74
Rosaceae	Prunus sp.	33
Rosaceae	Prunus spinosa L	41
Rosaceae	Prunus vachuschtii Bregaze	20
Rosaceae	Prunus vulgaris Mill	4
Rosaceae	Prunus x domestica L	296
Rosaceae	Pyracantha coccinea M. Roem	3
Rosaceae	Pyrus caucasica Fed	232
Rosaceae	Pyrus communis L	628
Rosaceae	Rosa canina L	11
Rosaceae	Rosa pimpinellifolia Boiss	13
Rosaceae	Rosa sp.	140
Rosaceae	Rubus caesius L	27
Rosaceae	Rubus fruticosus L	104
Rosaceae	Rubus idaeus L	268
Rosaceae	Rubus saxatilis L	19
Rosaceae	Rubus sp.	60
Rosaceae	Sorbus aucuparia K. Koch	18
Rosaceae	Sorbus boissieri C.K. Schneid	2
Rosaceae	Sorbus caucasigena Kom	57
Rosaceae	Sorbus torminalis C.Crantz	20
Rubiaceae	Coffea arabica L	1
Russulaceae	Lactarius deliciosus (L. ex Fr.) S.F. Grey	31
Russulaceae	Lactarius piperatus (L.) Pers	27
Russulaceae	Lactifluus piperatus (L.) Roussel	18
Russulaceae	Lactifluus volemus (Fr.) Kuntze	14
Russulaceae	Russula adusta Pers. Fr	6
Russulaceae	Russula emetica (Schaeff.) Pers	6
Russulaceae	Russula rosea Pers	23
Russulaceae	Russula virescens (Schaeff.) Fr	2
Rutaceae	Citrus limon (L.) Burm. f	15
Rutaceae	Citrus recticulata Blanco	5
Rutaceae	Citrus sinensis Osbeck	8
Rutaceae	Citrus unshiu Marcov	4
Rutaceae	Citrus x paradisi Macfad	2
Salicaceae	Salix caprea L	1
Sapindaceae	Acer pseudoplatanus L	2
Smilacaceae	Smilax excelsa L	91
Solanaceae	Capsicum annuum L	204
Solanaceae	Capsicum annuum L. "Sweet Bulgarian"	100
Solanaceae	Lycopersicum esculentum L	316
Solanaceae	Physalis alkekengi L	7
Solanaceae	Solanum melogena L	63
Solanaceae	Solanum pseudocapsicum L	2
Solanaceae	Solanum tuberosum L	347
Sparassidaceae	Sparassis crispa Wulfen	6
Staphyleaceae	Staphylea colchica Steven	116
Strophariaceae	Hypholoma fasciculare (Huds.) P. Kumm	6
Suillaceae	Suillus granulatus (L.) Roussel	14
Suillaceae	Suillus luteus (L.) Roussel	17
Taxaceae	Taxus baccata L	12
Theaceae	Camelia sinensis L	2
Tricholomataceae	Lepista sordida (Schumach.) Singer	18
Tricholomataceae	Tricholoma aurantium (Schaeff.) Ricken	1
Tricholomataceae	Tricholoma portentosum (Fr.) Quél	17
Tropaeolaceae	Tropaeolum majus L	1
Ulmaceae	Ulmus glabra Huds	3
Unidentified	Unidentified species	153
Urticaceae	Urtica dioica L	289
Violaceae	Viola arvensis L	1
Violaceae	Viola sp.	41
Vitaceae	Vitis labrusca L	26
Vitaceae	Vitis sylvestris W. Bartram	2
Vitaceae	Vitis vinifera L	538
Zingiberaceae	Elatteria cardamomum (L.) Maton	4

Table 6.

Distribution of mentions in plant families between garden and wild plants

Families	Garden	Wild	Families	Garden	Wild
Actinidiaceae	28	0	Liliaceae	6	39
Adoxaceae	6	128	Linaceae	0	1
Agaricaceae	6	225	Lythraceae	19	13
Amanitaceae	0	16	Malvaceae	14	157
Amaranthaceae	497	350	Marasmiaceae	0	12
Amaryllidaceae	853	302	Melanthiaceae	0	5
Annonaceae	1	0	Moraceae	237	11
Apiaceae	1422	490	Morchellaceae	0	13
Araceae	10	19	Musaceae	3	0
Araliaceae	1	0	Myrtaceae	11	0
Asparagaceae	7	52	Oleaceae	0	7
Asteraceae	492	252	Onagraceae	0	1
Auriculariaceae	0	10	Onocleaceae	4	31
Bankeraceae	0	10	Orobanchaceae	0	5
Begoniaceae	10	0	Oxalidaceae	2	1
Berberidaceae	10	42	Papaveraceae	4	28
Betulaceae	81	127	Physalacriaceae	0	93
Boletaceae	0	21	Phytolaccaceae	0	12
Boraginaceae	2	20	Pinaceae	3	44
Brassicaceae	899	99	Plantaginaceae	1	2
Campanulaceae	1	110	Pleurotaceae	2	92
Cannabaceae	39	13	Pluteaceae	0	28
Cantharellaceae	0	36	Poaceae	609	9
Caprifoliaceae	0	3	Polygonaceae	29	385
Caryophyllaceae	7	50	Polypodiaceae	0	10
Clavariadelphaceae	0	5	Polyporaceae	0	9
Convolvulaceae	15	2	Portulacaceae	6	79
Cornaceae	22	117	Primulaceae	0	55
Cortinariaceae	0	1	Psathyrellaceae	0	24
Corylaceae	1	3	Ramariaceae	0	12
Crassulaceae	0	32	Ranunculaceae	5	22
Cucurbitaceae	662	3	Rhamnaceae	1	2
Cupressaceae	0	2	Rhododendraceae	1	120
Dipsacaceae	0	1	Rosaceae	2683	1249
Dryopteridaceae	0	35	Rubiaceae	1	0
Ebenaceae	53	10	Russulaceae	3	124
Elaeagnaceae	1	9	Rutaceae	34	0
Ericaceae	4	472	Salicaceae	0	1
Euphorbiaceae	1	0	Sapindaceae	0	2
Fabaceae	738	101	Smilacaceae	0	91
Fagaceae	11	128	Solanaceae	1020	19
Fistulinaceae	0	6	Sparassidaceae	0	6
Fungi	2	225	Staphyleaceae	29	87
Gentianaceae	0	1	Strophariaceae	0	6
Geraniaceae	0	13	Suillaceae	0	31
Gomphaceae	0	6	Taxaceae	0	12
Grossulariaceae	226	99	Theaceae	2	0
Guttiferae	1	11	Tricholomataceae	0	36
Hericiaceae	0	1	Tropaeolaceae	1	0
Indet	24	126	Ulmaceae	0	3
Iridaceae	9	0	Urticaceae	31	258
Juglandaceae	222	27	Violaceae	0	42
Lamiaceae	550	403	Vitaceae	553	8
Lauraceae	23	4	Zingiberaceae	4	0
Lepiotaceae	0	24

Fig. 2.

Principal component analysis comparing the usage of plants based on their specific parts (bark, branches, buds, bulb, cones, flowers, fruit, latex, leaves, resin, roots, seeds, shoots, silk, stem, timber, tuber, whole plant) used. Contribution represents how each family contributes to the overall dissimilarity between regions based on their distance on the ordination. Arrows represent the specific plant parts used, small dots the samples and larger dots the centroid of each region

Most plants (65%) were eaten without complicated preparation, either raw (55%), or fried/cooked (e.g., 8% that were fungi). A full 5% of all mentioned plant-uses were for pickles / lactofermented (often stems), and a full 18% of all use reports were for Phkhali (boiled herb pie, especially in spring), 4% were used as spices, and around 2% for the distillation of alcohol. All other use categories (35) had much fewer mentions.

The richness of plant families was 66 in garden versus 97 families of wild plants, respectively, and the difference was highly significant. Other diversity indices also unequivocally pointed to a considerably more diverse family composition of wild versus garden plants as the differences between all the tested diversity indices appeared to be highly significant (Table 3).

Table 3.

Plant family diversity assessed by various indices

Index	Garden	Wild	P-value
Dominance, D	0.096	0.053	0.0001
Shannon H	2.709	3.525	0.0001
Evenness e^H/S	0.227	0.346	0.0001
Simpson index, 1—D	0.904	0.947	0.0001
Equitability J	0.647	0.769	0.0001
Fisher alpha	9.168	15.9	0.0001
Berger–Parker, BP	0.219	0.166	0.0001

P-values are calculated using randomization tests (or Permutation test, software PAST 4.2)

The regions of Georgia could be divided into three groups by the similarity of garden food plants as can be seen on the nMDS ordination graph (Fig. 3). This ordination seems to be influenced on the presence of large markets: Adjara, Samegrelo, Guria, and Kakheti which are lowland regions with large cities are joined by minimum distance versus Tori, Zemo Svaneti, Khevsureti, Tusheti and Javakheti, which are the most remote places. Kvemo Svaneti, Lechkhumi, Meskheti, Kvemo Kartli, Zemo Imereti, Zemo and Kvemo Racha, Mtianeti are moderately remote from large markets. The grouping of the regions closer to large markets might however have another distinct reason: Adjara, Samegrelo, Guria, and Kakheti are also the climatically warmest regions in Georgia, with the longest growing seasons. This allows the production of food plants almost all year round, and greatly reduces the dependency on foraging wild species.

Fig. 3.

nMDS ordination of regions by garden food plant species composition

For comparison, we assessed the usage of plants between regions based on their family, genus, specific parts used (root, shoot, or both) used, reproductive stages used (vegetative, reproductive, or both) and their specific parts used (bark, branches, buds, bulb, cones, flowers, fruit, latex, leaves, resin, roots, seeds, shoots, silk, stem, timber, tuber, whole plant), but at regional level and within different altitudinal ranges through non-multidimensional scaling (NMDS) followed by permutational multivariate analysis of variance (PERMANOVA) with 999 permutations and Euclidian distance. The detailed results are given in Table 4 and Appendix Tables 7, 8, 9, 10 and 11.

Table 4.

Pairwise comparisons with FDR p-value adjustment method of the different variables evaluated (plant family, plant genus, system used, general plant parts used, specific plant parts used, the usage) between altitudinal ranges after significant PERMANOVA analysis (Table Permanova)

Plant family
	0–500	1001–1500	1501–2000	2001–2500
1001–1500	0.0013
1501–2000	0.0013	0.0013
2001–2500	0.0013	0.0013	0.0013
501–1000	0.0490	0.0044	0.0013	0.0013
Plant genus
	0–500	1001–1500	1501–2000	2001–2500
1001–1500	0.0011
1501–2000	0.0011	0.0011
2001–2500	0.0011	0.0011	0.0011
501–1000	0.0180	0.0011	0.0011	0.0011
General plant parts used
	0–500	1001–1500	1501–2000	2001–2500
1001–1500	0.0300
1501–2000	0.3550	0.0300
2001–2500	0.4144	0.0300	0.3550
501–1000	0.0420	0.6270	0.0833	0.0300
General plant parts used
	0–500	1001–1500	1501–2000	2001–2500
1001–1500	0.0017
1501–2000	0.0722	0.0017
2001–2500	0.0017	0.0017	0.0017
501–1000	0.0271	0.6840	0.0288	0.0017
Specific plant parts used
	0–500	1001–1500	1501–2000	2001–2500
1001–1500	0.0017
1501–2000	0.0025	0.0017
2001–2500	0.0017	0.0017	0.0017
501–1000	0.0222	0.6670	0.0025	0.0017
Usage
	0–500	1001–1500	1501–2000	2001–2500
1001–1500	0.0133
1501–2000	0.0050	0.0957
2001–2500	0.0050	0.0840	0.3020
501–1000	0.0450	0.2833	0.0917	0.1750

Analyses were based on Euclidean distance and 999 permutations

Table 7.

Pairwise comparisons with FDR p-value adjustment method of plant family usage between regions after significant PERMANOVA analysis (Table Permanova)

	Adjara	Guria	Javakheti Plateau	Kakheti	Khevsureti	Kvemo Kartli	Kvemo Racha	Kvemo Svaneti	Lechkhumi	Meskheti	Mtianeti	Samegrelo	Tori	Tusheti	Zemo Imereti	Zemo Racha
Guria	0.0019
Javakheti Plateau	0.0019	0.0031
Kakheti	0.0019	0.0019	0.0159
Khevsureti	0.0019	0.0019	0.0044	0.0019
Kvemo Kartli	0.0019	0.0072	0.0019	0.0370	0.0031
Kvemo Racha	0.0117	0.0362	0.0019	0.0019	0.0019	0.0019
Kvemo Svaneti	0.0209	0.0031	0.0019	0.0044	0.0019	0.0019	0.0044
Lechkhumi	0.0608	0.0031	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019
Meskheti	0.0209	0.0378	0.0019	0.0031	0.0031	0.0082	0.0159	0.0126	0.0019
Mtianeti	0.0290	0.1400	0.0019	0.0290	0.0044	0.0544	0.0209	0.0095	0.0019	0.1068
Samegrelo	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019
Tori	0.0107	0.0229	0.0019	0.0019	0.0019	0.0031	0.0117	0.0031	0.0019	0.0393	0.0107	0.0019
Tusheti	0.0019	0.0019	0.0031	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019	0.0019	0.0031	0.0019	0.0019
Zemo Imereti	0.0031	0.0685	0.0019	0.0290	0.0019	0.0058	0.0019	0.0019	0.0019	0.0126	0.0082	0.0019	0.0019	0.0019
Zemo Racha	0.0044	0.0710	0.0082	0.0229	0.0019	0.0386	0.0159	0.0019	0.0019	0.0117	0.0561	0.0019	0.0031	0.0126	0.0181
Zemo Svaneti	0.0299	0.0019	0.0019	0.0019	0.0019	0.0019	0.0058	0.0082	0.0031	0.0474	0.0181	0.0019	0.0209	0.0019	0.0031	0.0019

Analyses were based on Euclidean distance and 999 permutations

Table 8.

Pairwise comparisons with FDR p-value adjustment method of plant genus usage between regions after significant PERMANOVA analysis (Table Permanova)

	Adjara	Guria	Javakheti Plateau	Kakheti	Khevsureti	Kvemo Kartli	Kvemo Racha	Kvemo Svaneti	Lechkhumi	Meskheti	Mtianeti	Samegrelo	Tori	Tusheti	Zemo Imereti	Zemo Racha
Guria	0.0012
Javakheti Plateau	0.0012	0.0012
Kakheti	0.0012	0.0012	0.0012
Khevsureti	0.0012	0.0012	0.0012	0.0012
Kvemo Kartli	0.0012	0.0012	0.0012	0.0012	0.0012
Kvemo Racha	0.0012	0.0022	0.0012	0.0012	0.0012	0.0012
Kvemo Svaneti	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012
Lechkhumi	0.0012	0.0065	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012
Meskheti	0.0012	0.0022	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012
Mtianeti	0.0055	0.0670	0.0012	0.0153	0.0012	0.0022	0.0073	0.0073	0.0012	0.0264
Samegrelo	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012
Tori	0.0012	0.0022	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0022	0.0012	0.0012
Tusheti	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012
Zemo Imereti	0.0012	0.0073	0.0012	0.0022	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0083	0.0012	0.0012	0.0012
Zemo Racha	0.0033	0.0584	0.0012	0.0073	0.0012	0.0022	0.0065	0.0012	0.0012	0.0022	0.0103	0.0012	0.0012	0.0012	0.0033
Zemo Svaneti	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0012	0.0022	0.0012	0.0012	0.0012	0.0012	0.0012

Analyses were based on Euclidean distance and 999 permutations

Table 9.

Pairwise comparisons with FDR p-value adjustment method of different plant system used (root, shoot, or both) between regions after significant PERMANOVA analysis (Table Permanova)

	Adjara	Guria	Javakheti Plateau	Kakheti	Khevsureti	Kvemo Kartli	Kvemo Racha	Kvemo Svaneti	Lechkhumi	Meskheti	Mtianeti	Samegrelo	Tori	Tusheti	Zemo Imereti	Zemo Racha
Guria	0.0065
Javakheti Plateau	0.0187	0.0038
Kakheti	0.4754	0.0121	0.2596
Khevsureti	0.4093	0.0038	0.0121	0.5112
Kvemo Kartli	0.4093	0.0139	0.0865	0.9340	0.4054
Kvemo Racha	0.0038	0.1808	0.0038	0.0139	0.0038	0.0065
Kvemo Svaneti	0.5393	0.0038	0.0231	0.7329	0.5763	0.6930	0.0038
Lechkhumi	0.2596	0.2546	0.0038	0.1539	0.0744	0.0544	0.0252	0.0415
Meskheti	0.5393	0.1396	0.0038	0.3965	0.3660	0.1808	0.0065	0.2546	0.2343
Mtianeti	0.7807	0.2720	0.0038	0.5731	0.5139	0.4038	0.0691	0.4871	0.2629	0.6245
Samegrelo	0.0038	0.0038	0.0065	0.0209	0.0038	0.0038	0.0038	0.0038	0.0038	0.0038	0.0038
Tori	0.0038	0.5112	0.0038	0.0038	0.0038	0.0038	0.2343	0.0038	0.0139	0.0038	0.0358	0.0038
Tusheti	0.4054	0.0038	0.0647	0.7222	0.7025	0.6091	0.0038	0.7323	0.0375	0.2629	0.4559	0.0065	0.0038
Zemo Imereti	0.0774	0.7439	0.0038	0.0680	0.0139	0.0340	0.2125	0.0321	0.3001	0.1104	0.2510	0.0038	0.4334	0.0163
Zemo Racha	0.6609	0.4054	0.0038	0.5273	0.4054	0.4038	0.1247	0.4054	0.6800	0.6800	0.7444	0.0065	0.1060	0.4054	0.4054
Zemo Svaneti	0.3660	0.1060	0.0038	0.1554	0.1396	0.1168	0.0038	0.1248	0.7108	0.6622	0.6887	0.0038	0.0095	0.1168	0.2149	0.7807

Analyses were based on Euclidean distance and 999 permutations

Table 10.

Pairwise comparisons with FDR p-value adjustment method of different general plant parts used (vegetative, reproductive, or both) between regions after significant PERMANOVA analysis (Table Permanova). Analyses were based on Euclidean distance and 999 permutations

	Adjara	Guria	Javakheti Plateau	Kakheti	Khevsureti	Kvemo Kartli	Kvemo Racha	Kvemo Svaneti	Lechkhumi	Meskheti	Mtianeti	Samegrelo	Tori	Tusheti	Zemo Imereti	Zemo Racha
Guria	0.0020
Javakheti Plateau	0.0020	0.0054
Kakheti	0.0020	0.0086	0.4630
Khevsureti	0.0020	0.0115	0.0115	0.3372
Kvemo Kartli	0.0020	0.0071	0.6074	0.6437	0.1026
Kvemo Racha	0.0020	0.3166	0.0020	0.0020	0.0020	0.0020
Kvemo Svaneti	0.6074	0.0071	0.0020	0.0101	0.0020	0.0020	0.0020
Lechkhumi	0.0020	0.0054	0.0020	0.0020	0.0020	0.0020	0.0020	0.0020
Meskheti	0.0302	0.3671	0.0020	0.1709	0.1593	0.0158	0.0158	0.0517	0.0020
Mtianeti	0.0915	0.4792	0.0020	0.5124	0.6437	0.1560	0.0666	0.0915	0.0020	0.7760
Samegrelo	0.0020	0.0020	0.0020	0.0020	0.0020	0.0020	0.0020	0.0020	0.0020	0.0020	0.0020
Tori	0.0020	0.1593	0.0020	0.0020	0.0020	0.0020	0.4439	0.0020	0.0020	0.0020	0.0038	0.0020
Tusheti	0.0020	0.0038	0.1885	0.3411	0.0857	0.5533	0.0020	0.0020	0.0020	0.0130	0.1676	0.0020	0.0020
Zemo Imereti	0.0020	0.5440	0.0038	0.0783	0.0260	0.0558	0.0920	0.0020	0.0020	0.0915	0.1916	0.0020	0.0020	0.0508
Zemo Racha	0.0020	0.2997	0.0526	0.3309	0.0915	0.2964	0.0535	0.0054	0.0020	0.0581	0.1511	0.0020	0.0020	0.4792	0.3992
Zemo Svaneti	0.2802	0.0260	0.0020	0.0020	0.0020	0.0020	0.0086	0.1119	0.0020	0.0250	0.0645	0.0020	0.0101	0.0020	0.0038	0.0020

Table 11.

Pairwise comparisons with FDR p-value adjustment method of specific plant parts used (bark, branches, buds, bulb, cones, flowers, fruit, latex, leaves, resin, roots, seeds, shoots, silk, stem, timber, tuber, whole plant) between regions after significant PERMANOVA analysis (Table Permanova)

	Adjara	Guria	Javakheti Plateau	Kakheti	Khevsureti	Kvemo Kartli	Kvemo Racha	Kvemo Svaneti	Lechkhumi	Meskheti	Mtianeti	Samegrelo	Tori	Tusheti	Zemo Imereti	Zemo Racha
Guria	0.0018
Javakheti Plateau	0.0018	0.0018
Kakheti	0.0018	0.0018	0.0267
Khevsureti	0.0018	0.0018	0.0033	0.0697
Kvemo Kartli	0.0018	0.0033	0.0057	0.3999	0.0057
Kvemo Racha	0.0018	0.1692	0.0018	0.0018	0.0018	0.0018
Kvemo Svaneti	0.2045	0.0018	0.0018	0.0018	0.0018	0.0018	0.0018
Lechkhumi	0.0057	0.0057	0.0018	0.0018	0.0018	0.0018	0.0018	0.0046
Meskheti	0.0046	0.1608	0.0018	0.0603	0.0018	0.0018	0.0046	0.0173	0.0018
Mtianeti	0.0267	0.3522	0.0018	0.3078	0.0096	0.0057	0.0324	0.0537	0.0018	0.6410
Samegrelo	0.0018	0.0018	0.0018	0.0018	0.0018	0.0018	0.0018	0.0018	0.0018	0.0018	0.0018
Tori	0.0018	0.0355	0.0018	0.0018	0.0018	0.0018	0.1349	0.0018	0.0033	0.0033	0.0046	0.0018
Tusheti	0.0018	0.0018	0.0148	0.0633	0.0433	0.0714	0.0018	0.0018	0.0018	0.0018	0.0071	0.0018	0.0018
Zemo Imereti	0.0018	0.2145	0.0018	0.0870	0.0033	0.0109	0.0222	0.0018	0.0018	0.0222	0.1272	0.0018	0.0033	0.0046
Zemo Racha	0.0018	0.1711	0.0018	0.2492	0.0083	0.1305	0.0267	0.0018	0.0018	0.0324	0.0668	0.0018	0.0018	0.0413	0.2493
Zemo Svaneti	0.0083	0.0057	0.0018	0.0018	0.0018	0.0018	0.0018	0.0787	0.0046	0.0334	0.0668	0.0018	0.0018	0.0018	0.0018	0.0046

Analyses were based on Euclidean distance and 999 permutations

The regions varied strongly in their species richness, based on species used (Fig. 4). These differences also might reflect the remoteness from large markets and severity of local climate.

Fig. 4.

Rarefaction of species richness of the garden food plants across the regions

Relationships among the regions in the case of wild food plants show a different and clearer pattern (Fig. 5). Adjacent regions in particular cluster together (Kvemo Zemo Racha, and Zemo Imereti; Samegrelo, Guria, Adjara, Lechkhumi and Kvemo and Zemo Svaneti; Meskheti, Javakheti, Kvemo Kartli; Mtianeti, Kakheti, Khevsureti, Tusheti). Like in the case of the garden food plants, species diversity of the wild food plants mentioned varied strongly (Fig. 6). Climate and the need for of the use of wild food plants (especially in high altitude villages) play a role in this variation. As we already showed in various previous publications, language, cultural group, ethnicity, education, or gender of the participants had no impact on the main use of food plants, nor any other uses [41–50].

Fig. 5.

nMDS ordination of regions by wild food plant species composition

Fig. 6.

Rarefaction of species richness of the wild food plants across the regions

Pkhali and Pickles—emblematic foods of the Caucasus

Of all food preparations the use of plants as ingredient of boiled herb preparations (mostly as —gazapkhuli pkhali = Spring Pkhkali, as the first vitamin source after winter), and as lacto-fermented or vinegar-based pickles are probably the most emblematic ones in the Caucasus, given that almost 50% of all food mentions were for phkhali, and almost 12% for pickled plants, and 8% for teas.

While the overall distribution of families, genera and their uses were similar between regions, overall most species were used in Guria. However, the knowledge distribution was most uneven for these food categories (Fig. 7). The altitudinal range between 1001 and 1500 m, followed by 1501–2000 m were clearly predominant when it came to diversity of plants used as well as uses (Fig. 8). This very unequal distribution of the most important families/genera, as well as their respective uses is reflected in Fig. 9. The altitudinal differences do not necessarily indicate however that the respective species did not grow also at lower altitudes. They simply indicate that at lower altitudes the participants rather preferred other food plants, and due to a lack of necessity were not interested in wild harvesting greens.

Fig. 7.

Relationship between families, genera and usage within regions

Fig. 8.

Relationship between families, genera and usage within the altitudinal gradients

Fig. 9.

Absolute mention of families and genera by region and altitudinal distribution

Only 60% of participants reported making pickles / lactofermented preparations. Of these, over 16% each came from Zemo Imereti and Khevsureti, and 12% each from Zvemo Svaneti, the Javakheti-Plateau, and Guria. The first regions represent all high altitude—short growing season areas, where the population does need to preserve food for winter. Guria is relatively warm—but very wet and snow-rich, which also might explain the prevalence of pickles. No participants whatsoever from Adjara, Samegrelo (the most subtropical regions) and Mtianeti (close to the capital Tbilisi) reported making pickles. Unsurprising, Kakhetians were also not enthusiastic about this form of preparation, because Kahketi is also a region famous for its large agricultural production. In contrast, in Tori and Tusheti there are simply less products that can be pickled. Preferred species (of a total of 79) for pickles were mostly Amaranthaceae (Amaranthus, Chenopdium), Apiaceae (especially the stems of Anthriscus, Chaerophyllum and Heracleum were pickled, but also, stems of Conium maculatum), Amaryllidaceae (all Allium species), and Polygonaceae (Polygonum and Rumex). In addition, Aruncus vulgaris (Rosaceae), Stapyllea colchica (Staphyleaceae). All of these were more important as pickles than "traditional European style species (Cucumis sativus, Capsicum etc.). The fermentation of the ferns Mattheucia struthiopteris (Onocleaceae) and Dryopteris filix-mas (Dryopteridaceae) was similar to what we observed, e.g., in the Himalayas.

The participants clearly indicated that some plants (e.g., Conium maculatum, Dryopteris filix-mas, Galanthus sp., Narcissus sp.) needed careful preparations, due to possible toxicity. However, given that these species might have even higher toxicity in other regions, e.g., Central Europe, the authors decided to not elaborate any further on preparation methods, given that these might not be sufficient to remediate toxicity of the same species outside the Caucasus.

In case of Pkhali, over 93% of all participants—from all regions—reported to use such boiled herbs, normally in Spring. This was surprising, as we had expected much more limited use in the climatically favorable regions. Nevertheless, Zemo Imereti (19% of all Phkhali preparations), Tori and Kvemo Racha (16% each), Tusheti (15%) and Khevsureti (14%)—all mountain regions with long winters, stood out as the real "herb eater" areas. In contrast to the pickled species, essentially only young leaves were used for pkhali, with great emphasis on the same families indicated in pickles. (All pickled plant species were also used for phkhali.) The overall number of species fused or pkhali was however much higher (197). The elaboration of phkhali often involves many steps to reduce the toxicity of species used, and in most cases a wide variety of herbs are included in each preparation. Interesting examples for the use of toxic species included the leaves of Solanum tuberosum, Veratrum lobelianum and Viola sp. Solanum tuberosum leaves for example are regarded as toxic worldwide, but are being eaten in the Caucasus and Albania [48]. Veratrum album (closely related to Veratrum lobelianum, and growing especially in Europe, is highly toxic), and Viola sp. (although especially the flowers are widely used in gastronomy) contains toxic Saponins. In all cases careful preparation was mentioned to make these species palatable. The authors explicitly decided to not give any recipes, given that many of the species are widespread, and compound composition—and with it possible toxic effects—might vary across the distribution range, so that a preparation method that sufficiently reduces toxicity in the Caucasus might not necessary be applicable in other areas.

Discussion

The use of food plant in Georgia while varied showed distinct overlap with other studies. However, the number of food plant species used—both cultivated and foraged in this rather small territory—was far higher than in most published studies from either wider region or the Mediterranean and Eurasia. Of all species, 388 were wild/wild collected, although a few of them also occurred as weeds in gardens. Even when deducting the fungal species (95), the remaining 293 vascular plant species are a mostly a much higher number than found in any other study in the wider region [73–106] (73:148 species; 74:87 species; 75:41 species; 76:40 species; 77:276 species; 78:119 species; 79:84 species; 80:68 species; 81:30–100 species for different European regions; 82:112 species; 83:139 species; 84:49 species; 85:15 species (although focusing on weeds only); 86:78 species; 87:419 species for all of Spain; 88:36; 89:77 species; 90:40 species; 91:11 species; 92:48 species; 93:83 species; 94:105 species; 95:73 species; 96:47 species; 97:115 species; 98:67 species; 99:78 species; 100:79 species; 101:35 species; 102:52 species; 103:63 species; 104:80 species; 105:88 species; 106:51 species).

Interestingly, even studies conducted in pastoralist cultures well-known for their use of wild foraged plants for food, e.g., in relatively close-by Kurdistan [107, 108] (107:54 species; 108:65 species), and Turkey [109] with 74 species showed a much more limited use of plants for food, even when not considering the 20% of taxa found in Georgia that were fungi. In many areas of the same cultural space, e.g., Dagestan [110] with 24 species, Azerbaijan [111, 112] (111:72 species; 112:73 species) and Amenia [113] with 66 species) the use of wild plants for food has been shown as in steep decline, although a strong signature of food plant use could still be found in markets of the Armenian capital Yerevan [114] with 148 species.

Outside the region, e.g., in China, it has been shown that typical agricultural communities use a very large number of wild species [115–117] (115: 185 vascular plant species and 17 fungal folk taxa; 116: 224 species; 117: 168 species). In many cases, however, wild plant use fell far short from the species numbers found in the Caucasus, e.g., [118–120] (118: 81 species; 119: 59 species; 120: 54 vascular plant species and 22 fungi).

The use of food species was not closely related to different vegetation zones in Georgia. This is a specific feature of food plants and differs from the use of plants in other categories, as has been previously shown [38–50].

The large number of species used in comparison with other areas confirmed our first hypothesis that given the long tradition of plant use, and the isolation under Soviet rule, plant use both based on home gardens and wild harvesting would be more pronounced in Georgia than in the wider region. In addition, the very large number of wild vegetables in Georgia might underline the hypothesis that the use of such wild "greens" is a byproduct of the Neolithic revolution, given that the region is indeed a cradle of agriculture as indicated previously [9, 13, 14].

We found a rather widespread use of foodplants across Georgia, which can partly be explained by mixture of populations from varied regions through migration and Soviet population moves, which also confirmed our hypothesis that food plant use knowledge would be widely and equally spread in most of Georgia.

Finally, we indeed found that in the very fertile agricultural regions in Eastern (Kakheti) and Western (lower Ajara, Samegrelo) Georgia, plant use knowledge was indeed more limited. However, this does not explicitly confirm our third hypothesis that such regions would show knowledge loss, as the limited use of plants may already have persisted a long time, and historic comparative data are not available.

Conclusions

This study reported on 535 plant and fungal taxa used in Georgia as food. As many mountain regions all over the world, the rural areas of the Georgian Caucasus have suffered a constant population decline for decades, due to harsh economic conditions and lack of modern infrastructure [1, 24, 121–124]. While this has greatly accelerated the loss of traditional agricultural practices, it seems to have affected the use of wild gathered food plants as well as species grown in home gardens to a much more limited extent in Georgia. The home gardens in Georgia clearly continue serving as socio-ecological memory, and an irreplaceable part of Georgian culture, rather than the widely growing popularity of gardening and foraging found all over Europe [125]. The great variety of food plant species used in the Georgian Caucasus provides a reservoir for food security for the region, as well as a source of important food plant germplasm for international agriculture. This greatly underlines the importance of Georgia as an ancient center of crop domestication and diversification, making Georgia clearly one of the most diverse food plant cultures in wider Eurasia, and the center of what we would like to coin as "Caucasus—Asia Minor—Balkans cultural complex."

Appendix

See Tables 5, 6, 7, 8, 9, 10 and 11.

Authors' contributions

RWB, NYPZ, SS, ZK, DK, MK, DT, and KB designed the study; RWB, NYPZ, SS, ZK, DT, MK, and KB conducted the fieldwork, ZK and IUR conducted the main statistical analysis; RBU, NYPZ, and ZK analyzed the data and wrote the manuscript; all authors read, corrected and approved the manuscript.

Funding

This study was funded through Saving Knowledge funds. The funding body itself has no direct role in the design of the study, collection or analysis of the data and use of results.

Availability of data and materials

The anonymized raw data are deposited under Open Science Network: https://osf.io/9kdtw/?view_only=93a8748c003f4770bc4a2bb332647429

Declarations

Ethics statement

Before conducting interviews, prior informed consent was obtained from all participants. No further permits or ethics approval were required.

Consent for publication

This manuscript does not contain any individual person's data, and further consent for publication is not required.

Competing interests

The authors declare that they have no competing financial interest.