From Biodiversity to Musketry: Detection of Plant Diversity in Pre-Industrial Peloponnese during the Flora Graeca Expedition

As the interest in natural, sustainable ecosystems arises in many fields, wild plant diversity is reconsidered. The present study is based on extant literature evidence from the journey of John Sibthorp (Professor of Botany, Oxford University) to Peloponnese (Greece) in pre-industrial time. In the year 1795, Peloponnese was a botanically unknown region, very dangerous for travellers and under civil unrest, in conjuncture with a pre-rebellion period. Our study reveals approximately 200 wild plant taxa that were collected from Peloponnese localities in 1795, transported to Oxford University (UK), and quoted in the magnificent edition Flora Graeca Sibthorpiana of the 19th century. Moreover, these plants currently constitute a living collection in Peloponnese, confirmed according to updated data on the vascular Flora of Greece. The presented lists constitute a source of information for plant biologists, linking the past to the present, shedding light on the study of adaptive traits of wild Mediterranean plants and revealing the temporal dimension of natural history. Nowadays, increasing and thorough understanding of the considered plants’ functionality to abiotic and biotic environmental stimuli provides a new framework of sustainability and management options.


Introduction
In the 18th century, travelers' journey to Greece was also a journey through its history. The naturalists' travels were explorations, linked to searching for specimens of natural history. The travelers' observations became a way of identifying and revealing cultural and economic changes that have occurred over the last centuries. The botanical expeditions and the collections of specimens connected observations and descriptions with landscapes and environmental conditions; plants had been there for thousands of years, linked to the history and adapted to abiotic and biotic conditions of the localities [1][2][3][4][5].
John Sibthorp (1758Sibthorp ( -1796, Professor of Botany in the University of Oxford, decided to travel to unexplored areas of Greece, collecting and recording botanical specimens in the late 1780s and 1790s; at that time, Greece was an unknown region, very dangerous and difficult to visit owing to diseases, civil unrest, and bandit groups-known as armatoloi and klephts-that included illiterate peasants, artisans, and local clergy, together with the local notables and landowners in Peloponnese [6][7][8].
Sibthorp's main interest was linked to plants known since the classical antiquity and mainly quoted in the texts of Dioscorides (1st century AD) [9][10][11][12][13][14]. During the first exploration from 1786 to 1787, Sibthorp was accompanied by the Austrian painter Ferdinand Lucas Bauer (1760-1826) as his draughtsman [6,7]; this was a time when travelers were accompanied by a professional artist, whose work supplemented their discoveries with visual evidence [15][16][17][18]. Actually, the magnificent, illustrated edition Flora Graeca Sibthorpiana (hereafter FGS), published from 1806 to 1840, contains botanical hand-coloured engravings that are important icons of the Mediterranean flora [7,19,20]. John Sibthorp and his companion undertook a second botanical expedition to the Levant from 1794 to 1795. During this journey, they arrived in Peloponnese (Morea is

Materials and Methods
This research is based on our survey of written sources, i.e., books, travel reports, letters, diaries, plant catalogues, online published, and printed archives mainly linked to the "Flora Graeca" expedition in Peloponnese (Greece) in 1795 [6,7]. Two copies of FGS, i.e., a copy adorned the National Library of Greece since 1916 and another copy acquired by the Gennadius Library of Athens in 1967 were surveyed. Moreover, we studied the digitized published hand-coloured engravings and the original watercolours, together with the Mediterranean scenes that are freely available and accessible online via Digital Bodleian (https://digital.bodleian.ox.ac.uk/collections/flora-and-fauna-graeca/, accessed on 9 October 2022). In addition, rigorous research of the Florae Graecae Prodromus [30] (hereafter Prodromus) housed in the Department of Botany at National and Kapodistrian University of Athens in Greece was carried out; it has to be noted that the Prodromus contains indexes of modern Greek vernacular names of plants (Index Nominum Graecorum, pp. 383-391), ancient Greek names of plants quoted in Dioscorides' codex (Index Dioscoridem, pp. 392-404), and scientific names of plants (Index Generum et Synonymorum, pp. 405-422), as well as plant locality data [31]. Furthermore, two books were taken in consideration; the first by Robert Walpole (1781-1856, an English classical scholar with degrees from Trinity College at Cambridge in UK and Merton College at Oxford in UK, who travelled to Greece; his Memoirs including notes of various travelers' diaries, among them Sibthorp's and his companion [32] were first published in 1817) and the second by John Bacon Sawrey Morritt (1772-1843, who immediately after his BA degree from St. John's College at Cambridge in UK, started on the travels described in his book that was first published in 1914; Morritt travelled over a considerable part of Peloponnese in 1795 [33]). A plant taxon was included in the results if there was a record in Prodromus stating locality data from Peloponnese. Information linked to the currently accepted plant nomenclature and distribution was derived from the Flora of Greece web (https://portal.cybertaxonomy.org/flora-greece/, accessed on 21 October 2022).

Peloponnese Tour
In Figure 1, the Peloponnese tours followed by Sibthorp and Morritt in 1795 are depicted in red and green lines, respectively. Sibthorp and his colleagues travelled from the island of Zakynthos to the port of Skaffidia (Ileia County); their route included Pyrgos, Lalla, and Tripolis, passing through several villages. The tour continued to Palaiepiscopi, In Figure 1, the Peloponnese tours followed by Sibthorp and Morritt in 1795 are depicted in red and green lines, respectively. Sibthorp and his colleagues travelled from the island of Zakynthos to the port of Skaffidia (Ileia County); their route included Pyrgos, Lalla, and Tripolis, passing through several villages. The tour continued to Palaiepiscopi, ancient Tegea, and Arcadia. Next, they travelled to Argos and visited ancient Mycenae as well as Napoli di Roamin (Nafplion) in Argolida County. Then, they travelled to Korinthos and Patras, continued in Achaia County through villages, and proceeded to Ileia County again; from there, they followed different directions until they arrived in Kalamata (Messinia County). After Kalamata, they proceeded to Kutchuk Maina, Kardamili, Sparta (Laconia County), and Mystras; from there, they continued to Messini and Petallida and on 25 April 1795 they arrived at Zakynthos and, by ship, returned to England. Morritt's journey started from Kalamata; he visited Kutchukmaina, Palaeocastro and ancient Thuria (Messinia County), Corone, Abia, and Kitreés and, through various villages, went to Kardamili/Cardamyla; he arrived by boat at Platsa and then continued to Oetylos, Marathonisi (ancient Gythium), and Mystras (Laconia County).

Plant Diversity in Pre-Industrial Peloponnese
Our study provides evidence for 183 plant taxa grown in pre-industrial Peloponnese, which had been collected during Sibthorp's expedition, drawn and cited in FGS (Table 1). Moreover, 21 plants quoted in Prodromus and linked to localities of Peloponnese, but neither drawn nor cited in FGS, were found ( Table 2). Although citations for prickly pear [Opuntia ficus-indica (L.) Mill.], walnut (Juglans regia L.), and mulberries (Morus nigra L.) were found in the considered archival research concerning Peloponnese, these plants were

Plant Diversity in Pre-Industrial Peloponnese
Our study provides evidence for 183 plant taxa grown in pre-industrial Peloponnese, which had been collected during Sibthorp's expedition, drawn and cited in FGS (Table 1). Moreover, 21 plants quoted in Prodromus and linked to localities of Peloponnese, but neither drawn nor cited in FGS, were found ( Table 2). Although citations for prickly pear [Opuntia ficus-indica (L.) Mill.], walnut (Juglans regia L.), and mulberries (Morus nigra L.) were found in the considered archival research concerning Peloponnese, these plants were neither drawn nor cited in both FGS and Prodromus. It should be mentioned that the botanist Sir James Edward Smith (1759-1828)-founder and first president of the Linnean Society of London-wrote the texts for the plants attested in FGS and Prodromus and excluded all species he regarded as not being part of the natural flora.   In 1795, in western Peloponnese, Salicornia fruticosa L. was observed growing near lake banks, Asphodelus ramosus L. near rivers, and Bromus rubens L. in between cultivated fields. Stands of Phillyrea latifolia L., Erica arborea L., Arbutus unedo L., Pistacia lentiscus L., vernal (spring) Crocus flavus Weston, and primroses (Primula vulgaris Huds.) in bloom-observed in early March 1795-were encountered. In the southern Peloponnese (county of Messinia), black mulberry trees (Morus nigra L.) and prickly pear surrounded many villages. Moreover, they depicted fig trees (Ficus carica L.), grapevines, cotton, grains, corn, olive trees, Euphorbia exigua L., Euphorbia spinosa L., Lolium perenne L., and Orobanche ramosa L. Some regions produced flax and tobacco. In the eastern Peloponnese, Quercus species, as well as corn, grains, grapevines, olive trees, fig trees, mulberry trees, and chestnut trees, had been detected. In the central Peloponnese (county of Arcadia), they visited oaks' forest; moreover, they observed a huge walnut tree (Juglans regia L.), Hyacinthus romanus L., and Hyacinthus spicatus Sm. in bloom. In addition, the presence of floating crystal-wort (Riccia fluitans L.) and Boletus (a genus of mushroom-producing fungi that comprises over 100 species) and the use of truffle were mentioned. Cultivation of pear trees with open blossoms (10 March 1795) and corns grown among the remains of cities and temples of the ancient Greek territories were detected. John Sibthorp arrived in Peloponnese bearing a mode of seeing, endowing the professorship of "Agriculture and Rural Economy" in the University of Oxford, thus the state of the agriculture in Peloponnese attracted his attention in 1795; the cultivation of corn (Zea mays L.), cotton (Gossypium hirsutum L.), millet (Panicum repens L.), tobacco (Nicotiana tabacum L.), and wheat (Triticum junceum L. and Aegilops comosa Sm.) was detected.

Discussion
Professor John Sibthorp and his colleagues visited Greek territories twice in preindustrial time, i.e., 1786-1787 and 1794-1795, and collected wild plants grown under natural conditions [7,16,34]. It was an outstanding achievement, considering the duration, the collections of specimens of plants from which "a legacy of 2462 pressed specimens are still preserved in the Sibthorpian Herbarium" [35] (Figure 2), and the geographical coverage, during the above-mentioned botanical expeditions. Moreover, a number of specimens found in Kew are of considerable importance as supplementing Sibthorp's collection at Oxford [36]; these specimens have been published [36] according to the sequence of plants cited in Prodromus [30].

Discussion
Professor John Sibthorp and his colleagues visited Greek territories twice in pre-industrial time, i.e., 1786-1787 and 1794-1795, and collected wild plants grown under natural conditions [7,16,34]. It was an outstanding achievement, considering the duration, the collections of specimens of plants from which "a legacy of 2462 pressed specimens are still preserved in the Sibthorpian Herbarium" [35] (Figure 2), and the geographical coverage, during the above-mentioned botanical expeditions. Moreover, a number of specimens found in Kew are of considerable importance as supplementing Sibthorp's collection at Oxford [36]; these specimens have been published [36] according to the sequence of plants cited in Prodromus [30]. The revived interest in FGS is partially due to recent publications [22,28,[37][38][39], but mainly to biodiversity issues raised under the threat of climate change, which gives another dimension to the whole achievement. Moreover, exhibitions dedicated to the concept and the content of Flora Graeca Sibthorpiana contributed to public awareness, e.g., in Oxford entitled "Painting by numbers" (Bodleian Library, 29 -9 July 2017, https://treas-  The revived interest in FGS is partially due to recent publications [22,28,[37][38][39], but mainly to biodiversity issues raised under the threat of climate change, which gives another dimension to the whole achievement. Moreover, exhibitions dedicated to the concept and the content of Flora Graeca Sibthorpiana contributed to public awareness, e.g., in Oxford entitled "Painting by numbers" (Bodleian Library, 29 -9 July 2017, https://treasures. bodleian.ox.ac.uk/treasures/flora-graeca/ accessed on 9 May 2017) and Athens entitled "Flora Graeca" (Gennadius Library, 8 March-4 July 2016, https://www.ascsa.edu.gr/ events/details/flora-graeca-exhibition, accessed on 8 March 2016).
In Table 1, we compiled a list of 183 wild plants cited in FGS and located in Peloponnese, which is indicative of the biodiversity, environmental physiology, phenology, and short flowering season in response to drought conditions, i.e., during the period of spring rainfall and the concomitantly active pollinators [40,41]. The later generations of plant biologists studied plant species grown in geographic locations visited by Sibthorp and his companion in Peloponnese, increasing the overall knowledge about distribution, ecophysiology, and taxonomy of plants quoted in FGS and Prodromus [42][43][44][45][46][47][48][49]. The mediterranean-type climate is characterized by a marked seasonality, typified by the alternation of a hot and dry period with a cold and wet period. For example, Sibthorp observed open flowers of Anemone coronaria L., Oxalis corniculata L., and Asphodelus ramosus L. on 27 February 1795, as well as of Crocus flavus Weston in early spring (cited as Crocus aureus in FGS and Crocus vernus latifolius aureus in Prodromus, vol. I, pp. [24][25]; such observations are supported by recent publications [5,50,51]. Moreover, in the 21st century, it is known that seasonal blossom is related to adaptive floral traits; for example, the study of petals revealed a surface nano-sculpture that declines water droplet adhesion and enhances the water repellence of these fragile floral tissues, which are exposed to the rainy conditions of the early spring flowering season [52][53][54]. In Anemone coronaria L., the temperature plays a critical role in the onset of dormancy [55]. Other species possess deeply rooted systems that enhance drought resistance (e.g., Myrtus communis L., Pistacia lentiscus L., and Quercus species). In addition, recent research revealed leaf functional traits linked to hydrophobicity and water status, highlighting species' responses to drought conditions [56][57][58]; this may be critical for resilience in the face of increasing drought stress.
Sibthorp and his companion visited a mountainy area, barren and stony beyond conception; it was hard work botanizing under harsh field conditions. The earth, washed by the rains and torrents from the higher parts, was supported on a plethora of terraces cultivated with wheat, cotton, maize, and millet, while olives and mulberry trees seemed to grow out of the rocky substrate itself. However, carpets of geophytes and numerous annual plants produced a spring flowering distinctive to the human eye. The results from this tour in the late 1790s, in pre-industrial landscapes, barely resembled the area we see today in Peloponnese, and brought information about numerous unknown to science (those days) wild plants, oak woodlands, pine forests, crops, cultivated areas, and arable lands of the monasteries [62]. Nowadays, several places of Peloponnese that Sibthorp visited in 1795 are included in the European network Natura 2000-i.e., the cornerstone of European Union nature conservation policy-of designated sites (https://eunis.eea.europa.eu/sites, accessed on 18 October 2022) relevant for flora and habitat protection [63-65], e.g., mountainy landscapes such as Parnonas: GR2520006, Mainalo (Arcadia): GR2520001, and Taygetos: GR2550006, as well as Folois plateau: GR2330002 and Olympia: GR2330004. Other progression was also recorded; that is, information linked to the current distribution of the considered plants, confirmed via the Flora of Greece web, contributed to our knowledge about natural stands of wild plants.
According to our study, on one hand, among the plants found in Peloponnese in 1795 and cited in FGS and Prodromus, there are species either widely distributed or grown in restricted areas, e.g., Achillea taygetea Boiss. & Heldr., Erophaca baetica Boiss., Saxifraga sibthorpii Boiss., and Scilla messeniaca Boiss. On the other hand, Zea mays L., originated from the Americas and found among the few cultivated species in isolated valleys in Peloponnese in pre-industrial time, might be attributed to the Venetian occupation of Peloponnese (1688-1715); during that period, when the area was dependent on the European market, plants might have been a product of cross-cultural communication between the conquerors and conquered [66][67][68][69][70].
Sibthorp's expedition in Peloponnese contributed to our understanding of botany in the field and revealed the diversity of plants grown in their habitats, in pre-industrial time. Historical time was linked to a gradually known plant diversity, as locations were explored and knowledge about the natural fertility of the land increased. However, anthropogenic pressure maintained by human activities, grazing, and fires in Peloponnese added to environmental stresses and caused profound transformation in the natural landscape, reducing the distribution of indigenous plants and enhancing a widespread concern about the extent of habitat and species loss [71][72][73][74][75][76]. This means that whatever effort can be made to study, maintain, and protect the diversity of ecosystems in this region is closely connected to a sustainable future, via the preservation of numerous plant taxa cited in the monumental FGS and Prodromus. Nowadays, Oxford Botanic Garden in UK (where visitors can enjoy the full sensory experience of walking through an aromatic Mediterranean landscape while learning about the work of Sibthorp and Bauer and its important botanical and horticultural legacy [35]) and Diomedes' Botanic Garden in Greece (due to the fact that administration of Diomedes' Botanic Garden is directly linked to the staff of the National and Kapodistrian University of Athens in Greece, this Garden has also been used for relevant, educational programs [37]) contain living collections of Mediterranean plants cited in FGS, which may be perceived as celebrations for Flora Graeca expeditions and FGS [35,37]. However, a larger number of plants quoted in FGS and Prodromus may be introduced and cultivated in the above-mentioned botanic gardens and/or the network of botanic gardens in Greece, in order to detect the diversity and the life-cycle of wild plants within the context of the seasons, floral colours in Mediterranean ecosystems, and collection and deposition of seeds in seed-banks. As such, botanic gardens can be used as common gardens, where researchers can conduct unmatched comparative research studies of plant ecophysiology, morphology, anatomy, and responses to climate change [77,78]. It is worth mentioning that Sibthorp introduced new species into English horticulture; moreover, he returned to Oxford from his eastern Mediterranean explorations with seeds, bulbs, and corms for the Botanic Garden, but few details of these collections have survived, and the plants and any knowledge about their propagation have been lost through many routes [7] (p. 180) and neglected [79] (p. 102).
This work provides a novel and valuable insight into the development of early plant environmental biology and is an important element of timelessness aspects of botany [80,81]. The study of plant diversity in Peloponnese peninsula, during the pre-rebellion period in Greece, tracing long-term changes in the region, is also a reminder that nature is often a repository at which nations look when crafting their identity.

Conclusions
The interest in archival material has been revived on account of research for a biodiversity threatened by climatic change. In this context, our research gives prominence to approximately 200 wild plant taxa found in Peloponnese (Greece)-most of them quoted in the magnificent edition Flora Graeca Sibthorpiana of the 19th century-and few cultivated introduced plants, all grown under ambient conditions and exposed to environmental stresses of the eastern Mediterranean during the pre-rebellion period, representing plant environmental issues in pre-industrial time, which have not hitherto been published.