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Article

Archaeological Areas as Habitat Islands: Plant Diversity of Epidaurus UNESCO World Heritage Site (Greece)

by
Maria Panitsa
*,
Maria Tsakiri
,
Dimitra Kampiti
and
Maria Skotadi
Laboratory of Botany, Department of Biology, University of Patras, GR26504 Patras, Greece
*
Author to whom correspondence should be addressed.
Diversity 2024, 16(7), 403; https://doi.org/10.3390/d16070403
Submission received: 27 May 2024 / Revised: 26 June 2024 / Accepted: 8 July 2024 / Published: 12 July 2024
(This article belongs to the Special Issue 2024 Feature Papers by Diversity’s Editorial Board Members)
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Abstract

:
The Epidaurus UNESCO World Heritage site (EPD) is a famous archaeological area that is located in a small valley in the Peloponnese and receives more than 250,000 visitors annually. The study of the plant diversity of the site is in the framework of a continuous research project concerning archaeological areas of the Peloponnese and in the context of a project by the Ministry of Culture, Education and Religious Affairs of Greece that started during 2023 to study the biodiversity of the archaeological areas of Greece. The main aim of this study is the exploration and analysis of the plant species composition and diversity of the Epidaurus archaeological area, with an emphasis on endemic plants, on ruderal and alien taxa as well as on environmental and disturbance indicators and the cultural ecosystem services they provide. This study revealed a high species richness consisting of 446 plant taxa. Most of them are Mediterranean and widespread, ruderals and medium disturbance indicators, but there are also 12 Greek endemic taxa. The richest in the taxa families are Asteraceae, Fabaceae and Poaceae. Therophytes predominate in the total flora registered and hemicryptophytes predominate in the endemics. Comparisons of the EPD’s plant diversity with other archaeological areas of Greece and the Mediterranean revealed its richness and unique character. Management and protection in archaeological areas such as the Epidaurus must focus on the sustainable conservation of their relationship with their natural environment.

1. Introduction

Archaeological areas and biodiversity influence and shape each other [1]. Archaeological sites are habitat islands and can be considered as repositories of valuable historical and cultural information and important havens for biodiversity, preserving the remnants of landscapes and ecosystems and providing valuable insights into human–environment interactions in the past [2,3,4,5,6,7,8]. Archaeological areas often become hotspots for biodiversity and refugia for plant species, insects and birds under sustainable management [9,10]. Incorporating archaeological knowledge in ecosystem services (ESs) would provide a basis for understanding human influences on the environment and sustainable natural resource management [11].
Effective conservation of cultural and natural heritage and sustainable development are among the goals and criteria of the World Heritage Convention. Criterion IX concerns “on-going ecological and biological processes in the evolution and development of ecosystems and communities” and Criterion X concerns “the most important and significant natural habitats for in-situ conservation of biological diversity, including those containing threatened species of outstanding universal value from the point of view of science or conservation” (https://whc.unesco.org/en/criteria/, assessed on 30 April 2024).
The ancient sanctuary of Asclepius at Epidaurus, a major healing center in ancient Greece dedicated to the god of medicine, Asclepius, is one of the 19 UNESCO World Heritage Sites in Greece, and is famous for its 4th century BC well-preserved theater, one of the best-preserved ancient theaters in Greece, known for its excellent acoustics and which is still used for performances today (https://whc.unesco.org/en/list/491, assessed on 30 April 2024). The Epidaurus archaeological area receives more than 250.000 visitors every year, and among the long-term goals is that the site, with respect to the monuments, will provide a natural, cultural and archaeological park with high-level visitor services.
With respect to their natural environment and biodiversity, cultural landscapes, including archaeological areas, are poorly inventoried [12] and evaluated, even in protected natural areas [5,8,9,13,14,15,16,17,18,19,20]. Studies concerning plant species diversity and communities of archaeological areas are limited (see, among others, [5,8,9,13,14,15,16,17,18,19,20,21,22,23,24]) and some of them focus on their coexistence with ancient structures (see, among others, [5,25,26,27,28,29,30,31,32,33,34]).
The study of the plant diversity of the UNESCO World Heritage site of Epidaurus started in the framework of a research project concerning archaeological areas of the Peloponnese [8]. The main aims of this study are as follows: a. the exploration and analysis of the plant species composition and diversity of the archaeological area of Epidaurus (Figure 1), with emphasis on endemic plants, on ruderal and alien taxa as well as on environmental and disturbance indicators and the cultural ecosystem services they provide; b. comparisons with other archaeological areas in Greece and the Mediterranean concerning their plant diversity in order to emphasize the importance of the UNESCO World Heritage site of Epidaurus as an important area in terms of its biodiversity conservation.

2. Materials and Methods

2.1. Study Area

The Epidaurus archaeological area (Figure 1) was inscribed in the UNESCO World Heritage sites in 1988 (https://whc.unesco.org/en/list/491, assessed on 30 April 2024). It is located in a small valley in the Peloponnese (Prefecture of Argolis) and covers a core area of 1393.8 ha and a buffer zone of 3386.4 ha. The area belongs to the thermo-Mediterranean vegetation zone and is characterized by xerothermic climatic conditions. Concerning the geology of the area, ammonites, like petrified snakes, in pink Triassic limestones at the entrance to the sacred precinct of Asclepios in Epidaurus may have attracted attention to that site of ancient healing with its emblem—the snake-encoiled caduceus [35].
The site is one of the most complete ancient Greek sanctuaries of antiquity, and comprises a series of ancient monuments spread over two terraces and surrounded by a preserved natural landscape. The theatre dates from the fourth century BC; it is one of the purest masterpieces of Greek architecture and the most authentic among the known theatres of the ancient world. The temple of Asklepios (the god of medicine) and the Tholos are also among the principal monuments, together with hospital buildings and temples that were devoted to healing gods during Greek and Roman times. The Epidaurus archaeological area is a natural, cultural and archaeological park with high-level visitor services and receives more than 250.000 visitors annually (https://whc.unesco.org/en/list/491, assessed on 30 April 2024).

2.2. Data Collection, Database, Floristic Analysis, Environmental and Disturbance Indicators and Ecosystem Services

In the present study, floristic data for the Epidaurus archaeological area (EPAA) are based on the collections and field observations of the authors during 2015 and 2021–2023. The sampling scheme used included fieldwork (plant specimen collections and observations) all over the studied area during all seasons of the year, focusing also on the floristic composition in distinct parts of the EPAA. For the determination of the plant material, refs. [36,37,38,39] were used. The families, genera, species and subspecies are listed within the major taxonomic groups in alphabetical order. Nomenclature, life forms, chorology, status categories and habitat preferences follow [40,41,42]; for the analysis, they have been grouped into the following five main chorological elements: widespread, Mediterranean, Balkan, Greek endemics and one group of chorological categories representing alien taxa of various origins, hereafter referred to as “alien” (established aliens are included in the floristic catalogue but casual aliens are not).
In the framework of the authors’ research, a database has been created including the plant taxa, their distribution in the studied area, their biological and chorological types and their habitat preferences according to the data provided by the “Vascular Plants of Greece” [40,41].
Ellenberg-type indicator values express plant ecological optima in the main environmental gradients [43,44]. The indicator values for light (L), temperature (T), moisture (F), soil reaction (R) and nutrients (N) follow [44,45]. For this study, indicator values for L are expressed on a scale from 1 to 9 (1: deep-shade plant; 9: full-light plant in fully irradiated places); for T (1: plants of cool sites, mean annual temperature < 8.5 °C; 9: plants of hot sites, mean annual temperature c. 20 °C and more); for F (1: strong drought indicator; 9: wetness indicator); R = soil reaction (1: indicator of strong acidity; 9: base and lime indicator); for N (1: indicator of nutrient-poorer sites; 9: indicator of very nutrient-rich sites); x = taxon species are indifferent concerning the parameter.
The indicator values for the disturbance frequency (DF), disturbance frequency—herb layer (DFH), disturbance severity (DS), disturbance severity—herb layer (DSH), for soil disturbance (SD) and for mowing frequency (MF) follow [46], as well as the European vegetation and flora data (https://floraveg.eu/, assessed on 30 April 2024). The indicator values for DF and DFH concerns the log10 mean inverse of the return time (in centuries) of all possible types of disturbance, including anthropogenic and natural disturbance, which is the mean interval between successive disturbance events (anthropogenic and natural disturbance); for DS and DSH, as a continuous value ranging from 0 (no change in biomass) to 1 (complete loss of plant cover); for MF, as the log10 mean inverse of the return time (in centuries) of disturbance, which is the mean interval between successive mowing events; for SD, as a continuous value ranging from 0 (no change in plant cover) to 1 (complete loss of plant cover), as described by [46]. The return time for disturbance is expressed in years and a 100-year value was assigned by default [46]. Separate values for DF, DS, DFH and DSH account for the fact that the DF and DS regimes in the tree and shrub layers differ in severity and frequency from the DFH and DSH regimes in the herb layer of the same community [46].
Cultural ecosystem services (CESs) and relevant indicators for different facets of plant diversity, identified as taxonomic diversity, life forms, species richness, presence of iconic/endemic/endangered species, Ellenberg-type environmental indicators, habitat diversity, landscape naturalness, monitoring sites (by scientists) and flower viewing, based on the CICES system (Common International Classification of Ecosystem Services), follow [47,48] and are in accordance with ecosystem services linked with different facets of biodiversity, as described by [49].

2.3. Comparisons with Other Archaeological Areas in Greece and the Mediterranean

Comparisons concerning plant diversity among the Epidaurus and other UNESCO World Heritage sites, or not, archaeological areas of the same phytogeographical area, namely, the Peloponnese, of Greece and of other Mediterranean countries are necessary in order to investigate the status of the plant species richness and diversity of the studied site using relative available data [8,22]. Different aspects of plant diversity are compared among the Epidaurus (EPD) and other archaeological sites, as follows: (i) A comparison with three other archaeological sites of the Peloponnese, namely, Akrokorinthos (AKK), Akronafplia (AKN) and Monemvasia (MON). Floristic data for AKK, AKN and MON are available from [8]. Total plant taxa lists are used (presence–absence of taxa per site) for the principal component analysis and hierarchical clustering. (ii) A comparison of ten archaeological sites in Greece, including AKK, AKN, MON and seven more studied by [22], namely, the Ancient Agora of Athens (AAA), Kolona Aegina (KA), Ancient Messene (AM), Nekromanteion of Acheron (NA), Ancient Forum of Thessaloniki (AFT) and Early Christian Amfipolis (ECA). In order to check for the presence or absence of the taxa registered more frequently on the 7 sites studied by [22] in the EPD and in the AKK, AKN and MON, lists of these plant taxa are compared with the one from the EPD using principal component analysis and hierarchical clustering. (iii) A comparison with two Italian areas, the archaeological park of the Neapolis of Syracuse (NEA) and the UNESCO World Heritage site of the Etruscan Necropolis of Tarquinia (TAR) [23,24,50], for which total plant taxa lists are available. The total number of plant taxa registered in each, the EPD, AKK, AKN, MON, NEA and TAR, and the surface area of these sites are used for investigating the species–area relationship.

3. Results

3.1. Floristic Composition and Local Distribution of the Taxa

The plant list includes 446 taxa (351 species and 95 subspecies), out of which 4 are Pteridophytes, 4 are Gymnosperms and 438 are Angiosperms, all belonging to 81 families and 303 genera. The floristic composition of the studied area is presented in the Supplementary Materials. Asteraceae (43 genera and 58 taxa), Fabaceae (24 genera and 62 taxa), Poaceae (29 genera and 40 taxa), Brassicaceae (17 genera and 19 taxa), Caryophyllaceae (12 genera and 19 taxa), Apiaceae (19 genera 21 taxa) and Lamiaceae (15 genera and 20 taxa) are the richest families. A total of 52.5% of the genera and 53.5% of the taxa of the total flora registered belong to these families. The genera richest in taxa are as follows: Medicago (11 taxa), Trifolium (13 taxa), Bromus (7 taxa), Vicia (6 taxa), Euphorbia (6 taxa), Geranium (6 taxa), Silene (6 taxa) and Plantago (5 taxa). Therophytes predominate (52.8%), followed by hemicryptophytes (18.8%), phanerophytes (13%), geophytes (10%) and chamaephytes (5.4%) (Figure 2).
Chorological analysis of the plant taxa registered showed that Mediterranean taxa predominate (53.9%), followed by widespread taxa (38.1%) (Figure 2). Twelve taxa are Greek endemics (2.7% of the plant taxa registered), and nine of them are range-restricted, belonging to six families and 11 genera. These include the following: Alkanna methanaea Hausskn., Anchusella variegata (L.) Bigazzi & al., Campanula andrewsii A. DC. subsp. andrewsii, Campanula drabifolia Sm., Centaurea raphanina subsp. mixta (DC.) Runemark, Chondrilla ramosissima Sm., Conium divaricatum Boiss. & Orph., Crocus laevigatus Bory & Chaub., Erysimum graecum Boiss. & Heldr., Onopordum argolicum Boiss., Scorzonera crocifolia Sm. and Stachys swainsonii subsp. argolica (Boiss.) Phitos & Damboldt. Hemicryptophytes dominate (66.6%) among the Greek endemics, followed by therophytes (16.7%), chamaephytes (8.3%) and geophytes (8.3%). Five taxa include Balkan endemics (1.1% of the plant taxa registered), namely, Astragalus suberosus subsp. haarbachii (Boiss.) V.A. Matthews, Bellevalia hyacinthoides (Bertol.) K.M. Perss. & Wendelbo, Crataegus heldreichii Boiss., Petrorhagia glumacea (Bory & Chaub.) P.W. Ball & Heywood and Verbascum undulatum Lam.
Seventeen taxa are alien-established (3.8%), belonging to 15 families and 16 genera. These include the following: Agave americana L., Amaranthus deflexus L., A. viridis L., Carpobrotus edulis (L.) N.E. Br., Cymbalaria muralis G. Gaertn. & al. subsp. muralis, Erigeron bonariensis L., Euphorbia prostrata Aiton, Iris albicans Lange, Lantana camara L., Lonicera japonica Thunb., Mirabilis jalapa L., Morus nigra L., Oxalis pes-caprae L., Prunus dulcis (Mill.) D.A. Webb, Punica granatum L., Robinia pseudacacia L. and Xanthium spinosum L. A total of 41.2% of these alien taxa are phanerophytes and 23.5% are hemicryptophytes, followed by therophytes (17.6%), geophytes (11.7%) and chamaephytes (5.9%). Alien taxa registered but not included in the plant lists, since they are not established, include the following: Jasminum mesnyi Hance, Lavandula angustifolia L., Melia azedarach Blanco and Populus × canadensis Moench.
Concerning the habitats they prefer, most of the taxa are ruderals (265 taxa, 59% of all taxa registered) belonging to 47 families and 185 genera. A total of 135 taxa (51% of the ruderals, 30% of all taxa registered) are exclusively found in ruderal areas and 130 are non-exclusive ruderals (Figure 3). Asteraceae (33 genera and 41 taxa), Fabaceae (17 genera and 34 taxa), Poaceae (18 genera and 27 taxa), Brassicaceae (14 genera and 16 taxa), Apiaceae (15 genera and 16 taxa) and Caryophyllaceae (8 genera and 12 taxa) are the richest families for ruderals, representing 66.7% of all the taxa registered belonging to these families (Figure 4). Taxa registered belonging to the family of Chenopodiaceae are ruderals, and those belonging to the families of Agavaceae, Aizoaceae, Amaranthaceae, Oxalidaceae and Punicaceae are all ruderals and alien.
Almost 29% of the taxa registered prefer xeric Mediterranean phrygana and grasslands exclusively (13.8%) or not (Figure 3). Two of the Greek endemics and range-restricted taxa, Campanula andrewsii subsp. andrewsii and Stachys swainsonii subsp. argolica, are chasmophytes exclusively preferring cliffs, rocks and walls.
The environmental preferences of the plant taxa registered are presented in Figure 5. Most of the taxa prefer open and not shady areas, full sunlight, hot and dry to slightly humid areas and rather basic soils rich in nutrients. A total of 23% of the taxa are indifferent to the “temperature” parameter and 12.5% are indifferent to the “soil reaction” parameter.
Figure 6 presents plant indicator values for DF, DFH, DS, DSH, MF and SD. Most of the plant taxa are indicators of low-to-medium DF and SD, high DS and also rather low indicators of MF. DFH shows higher values than DF for the majority of plants consisting of the herb layer, although DSH has low-to-medium values, lower than those for DS. Figure 7 presents plant indicator values for DF, DFH, DS, DSH, MF and SD for different life forms. Most of the therophytes are indicators of medium DFH and DSH, and low-to-medium MF and SD, while chamaephytes, geophytes and hemicryptophytes are high indicators of DFH, low-to-medium indicators of DSH and low indicators of MF and SD. Phanerophytes are indicators of high DS and of low DF, MF and SD.
Cultural ecosystem services (CESs) and relevant indicators for different facets of plant diversity, identified as taxonomic diversity, life forms, species richness, presence of iconic/endangered species (such as endemics), Ellenberg-type environmental indicators, habitat diversity, landscape naturalness, monitoring sites (by scientists) and flower viewing based on the CICES system (Common International Classification of Ecosystem Services), are presented in Table 1.

3.2. Comparisons with Other Archaeological Areas in Greece and the Mediterranean

Plant diversity of the archaeological areas of the Epidaurus differs significantly from those of three other archaeological areas (AKK, AKN and MON) of the same phytogeographical area, namely, the Peloponnese. Figure 8a presents the results of the principal component analysis and Figure 8b presents the results of hierarchical clustering, highlighting the dissimilarity of the Epidaurus.
Concerning the most common plant taxa among the archaeological areas of the EPD and ten other archaeological areas of Greece (AKK, AKN, MON, AAA, AFT, AM, AO, ECA, KA and NA), it is obvious that the EPD presents significant differences from the other areas. Figure S1 presents the results of the hierarchical clustering of the common plant taxa among the EPD and these 10 archaeological areas, based on the most common taxa of AAA, AFT, AM, AO, ECA, KA and NA as found by [22]. A total of 32% of these taxa were not registered in the EPD. Figure 9a presents the results of the principal component analysis and Figure 9b presents the results of the hierarchical clustering analysis, highlighting the dissimilarity of the EPD from the 10 other archaeological areas of Greece concerning the common plant taxa. The most common among these taxa have been registered in more than 50% of the areas, namely, Asplenium ceterach, Calendula arvensis, Capsella bursa-pastoris, Cynodon dactylon, Catapodium rigidum, Erodium malacoides, Gallium aparine, Geranium molle, Hyparrhenia hirta, Mercurialis annua, Papaver rhoeas, Plantago lanceolata, Reseda alba, Senecio vulgaris, Sonchus asper, Tragopogon porrifolius, Trifolium campestre, Umbilicus spp. and Veronica cymbalaria.
The surface areas of the EPD, AKK, AKN, MON and two Italian archaeological areas (NEA and TAR) explained 39.3% of the differences in their total plant species richness (R2 = 39.3%) (Figure 10).

4. Discussion

UNESCO’s World Heritage Committee defines cultural landscapes as geographical areas ‘representing the combined work of nature and man’. The role of cultural ecosystem services (CESs) with respect to what ecosystems can offer to people, such as aesthetic values, educational values or tourism and recreation possibilities, in heritage research can be estimated through different indicators [48]. The role that a UNESCO World Heritage site such as the Epidaurus archaeological area plays for people is much more than its historical value, to which a conservation scenario is mainly focused [51], and the role of community-based heritage management is more effective when people are emotionally attached to historic sites [52]. Cultural activities in archaeological areas are linked not only to heritage as an ecosystem service provided to users, but also with feelings of overall connections to both nature and culture [53,54,55]. An integrated estimation of the values of CESs, using social and ecological concepts, is necessary [56]. The relationship between heritage and natural components is a very important issue for the conservation and protection of the archaeological areas [57].
The Epidaurus UNESCO World Heritage site is characterized by the provision of several CESs and is famous for its cultural value. Among the CES indicators with physical/natural values, landscape naturalness and diversity, species richness and the presence of significant species are among the most common that are currently used, together with the number of environmental education-related facilities/events, the presence of vegetation, flower viewing, willingness to pay for environment improvements, protected natural areas, monitoring sites (by scientists), scenic beauty natural heritage and cultural sites [48,56,58].
The Epidaurus UNESCO World Heritage site is considered successfully managed concerning the ancient monuments and buildings (https://whc.unesco.org/en/list/491, accessed on 25 May 2024), receives a high number of visitors every year and, consequently, there are many and constant pressures and disturbances on its natural environment. Anthropogenic landscapes with a long history present a “not-natural” biodiversity, but rather form from the continuous interaction between man and nature over the time [59]. The species pool of an area reflects the long-term processes affecting it [60], and the composition of the plants associated with archaeological areas gives us the chance to understand the processes of species persistence on these habitat islands [61]. Archaeological areas, as with most cultural sites, are characterized by high species and community diversity, and their management, different than that of other areas, may increase them [2,3,4,5,6,8,9,62]. This is confirmed through the current study, which has revealed one more aspect of this famous area, its rich plant richness and diversity consisting mainly of Mediterranean and widespread plant taxa adapted to ruderal areas and frequent disturbances. The high representation of the richest taxa families of Asteraceae, Fabaceae and Poaceae, and also of therophytes, Mediterranean taxa and leguminous taxa (Trifolium, Medicago and Vicia are among the richest in the taxa genera), was expected [8,22,23,24,32,50,63] and expresses the combined effect of the environmental conditions of the Mediterranean and intense human impact (e.g., [64,65,66,67,68,69,70,71]). The functional traits of many plant taxa registered as dispersal mechanisms (zoochory for Poaceae and anemochory for Asteraceae) and the resistance to mowing for Fabaceae help them maintain their populations [13,22,28]. The rather low number of endemics can be explained by the simple topography of the studied area and the human impact. Hemicryptophytes and chamaephytes dominate among the endemics (75%) and are mainly found on the ancient walls and ruins exposed to extreme conditions (strong temperature variations, lack of water and soil, etc.) [3,8]. Other cultural areas of the Peloponnese presented higher proportions of endemics due to their more complex topography, including cliffs that hosts many endemics all over the Peloponnese and Greece [8,72].
Concerning alien taxa, they represent a low proportion of the total flora registered, and most of them prefer ruderal habitats. Phanerophytes predominate among the established alien taxa, and taxa such as Robinia pseudacacia, Punica granatum and Lantana camara, as well as the not-established Melia azedarach, are often present in archaeological areas [13,15,23,24,73,74]. Robinia pseudacacia, Carpobrotus edulis and Lantana camara are invasive species in Mediterranean ecosystems and among the 100 most dangerous species in Europe, strongly affecting native plant communities [75].
Disturbance is an important driver of plant biodiversity and function, and disturbance indicator values can be used to test how plant morphophysiology and the functional composition of plant communities respond along disturbance gradients [46,76]. Our results regarding disturbance (DF, DFH, DS, DSH, SD and MF) plant indicator values are in accordance with those of urban competitor and ruderal plants [77]. For example, according to [77], the dominance of annual plants, therophytes, relates to the general rapid life cycle of ruderal species and their abundance within urban indicators. Herbaceous hemicryptophytes and therophytes dominate in ruderal flora [8,70] and in the nitrophilous flora of archaeological areas adapted to periodic cutting and other management practices [5]. Additionally, Ellenberg indicator values that have only gentle differences among the different plant communities reflect the ecological pattern of the area studied [50].
The low similarity with other archaeological sites of the same phytogeographical area in Greece and Italy concerning the total flora, as well as the presence of the most common taxa registered in them, reflect the high plant species richness and diversity of the EPD. The surface area of the archaeological sites does not seem to be the only dominant factor affecting species richness, and since the topography of the EPD is rather gentle and the elevation is low, human impact could also be one of the main factors affecting, and increasing, its high plant diversity [78,79,80,81,82]. The majority of the taxa are therophytes, disturbance-tolerant and prefer ruderal habitats as most generalist plant taxa [2,4,5,6,70,77,82], but there are also specialists, endemic taxa that survive in places not strongly influenced by the large number of visitors and management practices. The presence of endemic taxa with conservation interest underlines the need for conservation strategies that consider the cultural and natural aspects of the studied area and for sustainable landscape management [6,13,50]. In summary, an in-depth knowledge of plant ecology could help to preserve the floristic richness of this site by applying focused management actions close to or on the monuments to avoid biodeterioration and mild actions in the remaining areas [12,22,27,28,31,33].
The Nature as Culture perspective confronts as a whole nature and human co-creations and underlines the significance of culture in “shaping and being shaped” by nature [83,84]. The evaluation and provision of guidelines for the sustainable management of cultural landscapes, as archaeological sites, assume knowledge and understanding of all aspects of biodiversity [5,6,13,85,86,87,88,89]. Archaeology and knowledge of the environmental settings provides economics and ecology with a long-term perspective on human–environment interactions and gives better chances for understanding the human impact on the environment and sustainable natural resource management [1].

5. Conclusions

Archaeological landscape conservation needs a comprehensive approach, including both cultural and natural aspects [5,6,8,28,29,86,87,88]. In the present study, the very rich plant diversity of the Epidaurus UNESCO World Heritage site, characterized as a habitat island, as with all archaeological areas, highlighted the need for scientific knowledge, not only of the historical elements but also of the biodiversity (plant species diversity) and their interaction so to produce the best management and conservation case-specific strategies.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/d16070403/s1, Table S1. Plant taxa registered in the Epidaurus UNESCO World Heritage site. Figure S1. Hierarchical clustering for the most common plant taxa species among the Epidaurus UNESCO World Heritage site and 10 more Greek archaeological areas, as found for 7 of them by [22].

Author Contributions

Conceptualization, M.P.; methodology, M.P.; validation, M.P., M.T., D.K. and M.S.; formal analysis, M.P. and M.T.; investigation, M.P., M.T., D.K. and M.S.; resources, M.P., M.T., D.K. and M.S.; data curation, M.T. and D.K.; writing—original draft preparation, M.P.; writing—review and editing, M.P., M.T., D.K. and M.S.; visualization, M.P. and M.T.; supervision, M.P.; project administration, M.P. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Data Availability Statement

Data are contained within the article and Supplementary Materials.

Acknowledgments

This research was supported in part by an ongoing research funded by the Natural Environment and Climate Change Agency (NECCA) of Greece, in the framework of the project "Biodiversity and Archaeological areas: Inventory of flora and fauna of archaeological areas". Fieldwork was carried out through special permits issued by the Ministry of Culture and Sports (Protocol Number: 70512-24/2/2023) and the Ministry of Environment and Energy (AΔA: ΨΥΓΡ4653Π8-ΦΕΡ), both supporting the project.

Conflicts of Interest

The authors declare no conflicts of interest.

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Figure 1. Studied area. Arrow on the left show location of the UNESCO World Heritage site of Epidaurus in Greece. Arrow on the right presents Ancient Theatre at the Asclepieion of Epidaurus.
Figure 1. Studied area. Arrow on the left show location of the UNESCO World Heritage site of Epidaurus in Greece. Arrow on the right presents Ancient Theatre at the Asclepieion of Epidaurus.
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Figure 2. Life form spectrum and chorological analysis of the total flora registered. Abbreviations: C = chamaephytes; G = geophytes; H = hemicryptophytes; P = phanerophytes; T = therophytes.
Figure 2. Life form spectrum and chorological analysis of the total flora registered. Abbreviations: C = chamaephytes; G = geophytes; H = hemicryptophytes; P = phanerophytes; T = therophytes.
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Figure 3. Habitat categories represented among all taxa. Abbreviations: R = agricultural and ruderal habitats; P = xeric Mediterranean phrygana and grasslands; W = woodlands and scrub; C = cliffs, rocks, walls, ravines and boulders; M = coastal habitats.
Figure 3. Habitat categories represented among all taxa. Abbreviations: R = agricultural and ruderal habitats; P = xeric Mediterranean phrygana and grasslands; W = woodlands and scrub; C = cliffs, rocks, walls, ravines and boulders; M = coastal habitats.
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Figure 4. Richest taxa families and representation of ruderal (blue bars) and non-ruderal (orange bars) taxa among them.
Figure 4. Richest taxa families and representation of ruderal (blue bars) and non-ruderal (orange bars) taxa among them.
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Figure 5. Preferences of the plant taxa registered to different environmental parameters. L = light (1: deep-shade plant; 9 full-light plant in fully irradiated places); T = temperature (1: plants of cool sites, mean annual temperature < 8.5 °C; 9: plants of hot sites, mean annual temperature c. 20 °C and more); F = moisture (from 1: strong drought indicator to 9: wetness indicator); R = soil reaction (from 1: indicator of strong acidity to 9: base and lime indicator); N = nutrient supply (from 1: indicator of nutrient-poorer sites to 9: indicator of very nutrient-rich sites); x = taxon species that are indifferent concerning the parameter (with the indicators as in [44,46]).
Figure 5. Preferences of the plant taxa registered to different environmental parameters. L = light (1: deep-shade plant; 9 full-light plant in fully irradiated places); T = temperature (1: plants of cool sites, mean annual temperature < 8.5 °C; 9: plants of hot sites, mean annual temperature c. 20 °C and more); F = moisture (from 1: strong drought indicator to 9: wetness indicator); R = soil reaction (from 1: indicator of strong acidity to 9: base and lime indicator); N = nutrient supply (from 1: indicator of nutrient-poorer sites to 9: indicator of very nutrient-rich sites); x = taxon species that are indifferent concerning the parameter (with the indicators as in [44,46]).
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Figure 6. Plant indicator values for DF, DFH, DS, DSH, MF and SD. Abbreviations: DF = disturbance frequency; DFH = disturbance frequency—herb layer; DS = disturbance severity; DSH = disturbance severity—herb layer; MF = mowing frequency; SD = soil disturbance.
Figure 6. Plant indicator values for DF, DFH, DS, DSH, MF and SD. Abbreviations: DF = disturbance frequency; DFH = disturbance frequency—herb layer; DS = disturbance severity; DSH = disturbance severity—herb layer; MF = mowing frequency; SD = soil disturbance.
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Figure 7. Plant indicator values for DF, DFH, DS, DSH, MF and SD of different life forms. Abbreviations: DF = disturbance frequency; DFH = disturbance frequency—herb layer; DS = disturbance severity; DSH = disturbance severity—herb layer; MF = mowing frequency; SD = soil disturbance; C = chamaephytes; G = geophytes; H = hemicryptophytes; P = phanerophytes; T = therophytes.
Figure 7. Plant indicator values for DF, DFH, DS, DSH, MF and SD of different life forms. Abbreviations: DF = disturbance frequency; DFH = disturbance frequency—herb layer; DS = disturbance severity; DSH = disturbance severity—herb layer; MF = mowing frequency; SD = soil disturbance; C = chamaephytes; G = geophytes; H = hemicryptophytes; P = phanerophytes; T = therophytes.
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Figure 8. Diagrams of (a) the principal component and (b) hierarchical clustering analyses concerning the plant diversity of the archaeological areas of the Epidaurus (EPD) and three other archaeological areas of the Peloponnese, including the Akrokorinthos (AKK), Akronafplia (AKN) and Monemvasia (MON).
Figure 8. Diagrams of (a) the principal component and (b) hierarchical clustering analyses concerning the plant diversity of the archaeological areas of the Epidaurus (EPD) and three other archaeological areas of the Peloponnese, including the Akrokorinthos (AKK), Akronafplia (AKN) and Monemvasia (MON).
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Figure 9. Diagrams of (a) the principal component and (b) hierarchical clustering analyses concerning the common plant taxa of the archaeological areas of the Epidaurus (EPD) and nine other archaeological areas of Greece, namely, the Akrokorinthos (AKK), Akronafplia (AKN), Monemvasia (MON), Ancient Agora of Athens (AAA), Kolona Aegina (KA), Ancient Messene (AM), Nekromanteion of Acheron (NA), Ancient Forum of Thessaloniki (AFT) and Early Christian Amfipolis (ECA).
Figure 9. Diagrams of (a) the principal component and (b) hierarchical clustering analyses concerning the common plant taxa of the archaeological areas of the Epidaurus (EPD) and nine other archaeological areas of Greece, namely, the Akrokorinthos (AKK), Akronafplia (AKN), Monemvasia (MON), Ancient Agora of Athens (AAA), Kolona Aegina (KA), Ancient Messene (AM), Nekromanteion of Acheron (NA), Ancient Forum of Thessaloniki (AFT) and Early Christian Amfipolis (ECA).
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Figure 10. Total plant species richness–area relationship of four Greek and two Italian archaeological areas (R2 = 0.393, p < 0.5). Abbreviations: A = area, S = number of species, Epidaurus (EPD), Akrokorinthos (AKK), Akronafplia (AKN) and Monemvasia (MON), the Neapolis of Syracuse, Sicily, Italy (NEA) and the UNESCO World Heritage site of the Etruscan Necropolis of Tarquinia, Italy (TAR).
Figure 10. Total plant species richness–area relationship of four Greek and two Italian archaeological areas (R2 = 0.393, p < 0.5). Abbreviations: A = area, S = number of species, Epidaurus (EPD), Akrokorinthos (AKK), Akronafplia (AKN) and Monemvasia (MON), the Neapolis of Syracuse, Sicily, Italy (NEA) and the UNESCO World Heritage site of the Etruscan Necropolis of Tarquinia, Italy (TAR).
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Table 1. Cultural ecosystem services (CESs) and relevant indicators according to [38,39] for different facets of the biodiversity identified based on the CICES system.
Table 1. Cultural ecosystem services (CESs) and relevant indicators according to [38,39] for different facets of the biodiversity identified based on the CICES system.
DivisionGroupClassV4.3 EquivalentCodeRelevant ES Indicators
Direct, in situ and outdoor interactions with living systems that depend on the presence in the environmental settingPhysical and experiential interactions with natural environmentCharacteristics of living systems that enable activities promoting health, recuperation or enjoyment through active or immersive interactionsExperiential use of plants, animals and land/seascapes in different environmental settings3.1.1.1T, L, S, E, EI, H, LN, M, F
Characteristics of living systems that enable activities promoting health, recuperation or enjoyment through passive or observational interactionsPhysical use of land/seascapes in different environmental settings3.1.1.2T, L, S, E, EI, H, LN, M, F
Intellectual and representative interactions with natural environmentCharacteristics of living systems that enable scientific investigation or the creation of traditional ecological knowledgeScientific3.1.2.1T, L, S, E, EI, H, LN, M,
Characteristics of living systems that enable education and trainingEducational3.1.2.2T, L, S, E, EI, H, LN, M, F
Characteristics of living systems that enable aesthetic experiencesAesthetic3.1.2.5T, L, S, E, H, LN, F
Elements of living systems used for entertainment or representationEntertainment3.1.2.4T, LN, F
T: taxonomic diversity; L: life form; S: species richness, E: presence of iconic/endangered species (no.); EI: Ellenberg-type indicator values; H: habitat diversity; LN: landscape naturalness; M: monitoring sites (by scientists); F: flower viewing.
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Panitsa, M.; Tsakiri, M.; Kampiti, D.; Skotadi, M. Archaeological Areas as Habitat Islands: Plant Diversity of Epidaurus UNESCO World Heritage Site (Greece). Diversity 2024, 16, 403. https://doi.org/10.3390/d16070403

AMA Style

Panitsa M, Tsakiri M, Kampiti D, Skotadi M. Archaeological Areas as Habitat Islands: Plant Diversity of Epidaurus UNESCO World Heritage Site (Greece). Diversity. 2024; 16(7):403. https://doi.org/10.3390/d16070403

Chicago/Turabian Style

Panitsa, Maria, Maria Tsakiri, Dimitra Kampiti, and Maria Skotadi. 2024. "Archaeological Areas as Habitat Islands: Plant Diversity of Epidaurus UNESCO World Heritage Site (Greece)" Diversity 16, no. 7: 403. https://doi.org/10.3390/d16070403

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