The beginnings of ecological thought in the Renaissance: an account based on the Libri picturati A. 18–30 collection of water-colours

Archives of natural history 34 (1): 87–108. 2007 © A. Zemanek, A. Ubrizsy Savoia & B. Zamanek 2007.

The beginnings of ecological thought in the Renaissance: an account based on the Libri picturati A. 18–30 collection of water-coloursALICJA ZEMANEKA, ANDREA UBRIZSY SAVOIAB and BOGDAN ZEMANEKC

A J. Dyakowska Botanical Museum and History of Botany Research Unit, Botanic Garden, Institute of Botany, Jagiellonian University, 31-501 Cracow, Kopernika 27, Poland (e-mail: [email protected]).B Botanic Garden, Department of Plant Biology, University ‘La Sapienza’ of Rome, Largo Cristina di Svezia 24,I-00165 Roma, Italy (e-mail: [email protected]).C Botanic Garden, Institute of Botany, Jagiellonian University, 31-501 Cracow, Kopernika 27, Poland (e-mail: [email protected]).

ABSTRACT: During the Renaissance ecological thinking emerged both in printed scientific works and in pictures showing plants against the background of their natural environment. A unique source for the history of plant ecology is the Libri picturati A. 18–30 collection of water-colours kept at the Jagiellonian Library in Cracow (Poland). This collection consists of 13 volumes of plant pictures, and contains about 1,800 images illustrating more than 1,000 taxa mainly from north-western Europe and the Mediterranean region, but also from Asia and America. Some of these pictures match with woodcuts in various works by famous Flemish botanists, mainly Charles de l’Écluse (Carolus Clusius) (1526–1609). Both the illustrations and their short annotations provide a synthetic review of the ecology of the Renaissance period. The paper deals with ecological issues which are found in the collection such as information on the climatic and edaphic requirements of some species, on plants occurring in various habitats and plant communities, plants representing principal growth forms, descriptions of particular adaptations to specific living conditions, for example the halophyte community of sea coasts or the parasitic flowering plants, and phenological observations. These trends can also be seen in printed publications of that time, and this collection mirrors them especially closely.

KEY WORDS: Renaissance botany – plant pictures – history of ecology – Charles de l’Écluse – Carolus Clusius.

INTRODUCTION

Knowledge of the relationships between plants and the environment, and of the seasonal changes in plants, has been, since the oldest times, formulated in close relation to agricultural and horticultural practices, but ecology was separated as a distinct branch of biological sciences only in the nineteenth century. The early precursors of ecological thinking were ancient scholars, particularly the founder of botany Theophrastus of Eresus (c. 370–285BC), who discussed, for example, the relationships of plants to the climate and soil condition, listed species peculiar to various types of habitat, and showed awareness of plant communities (Morton, 1981: 41; Greene, 1983: 1: 195–198). During the Renaissance, when the rediscovery of ancient knowledge was combined with new learning about the flora of Europe, as well as with exploring newly-discovered continents, ecological thinking emerged both in printed scientific treatises as well as in pictures showing plants against the background of their natural environment (Morton, 1981: 125–126; Piekiełko-Zemanek, 1986; Ubrizsy Savoia, 1998; Zemanek, 1998b: 32; Dobat, 2001: 16–17).

88 LIBRI PICTURATI AND THE BEGINNINGS OF ECOLOGICAL THOUGHT

Figure 1. Mandragora officinarum (30/85), one of the most famous ‘magic plants’. The annotation shows that its author did not support the old beliefs connected with this herb: “Grows in the mountains, abundantly in Gargano Mountains in Apulia; and it is sown and cultivated carefully in the gardens for its false miraculous properties. With us fruit ripens in July and August”. © Jagiellonian Library, Graphics Department, Cracow, Poland.

http://www.eupjournals.com/action/showImage?doi=10.3366/anh.2007.34.1.87&iName=master.img-000.jpg&w=342&h=520

89LIBRI PICTURATI AND THE BEGINNINGS OF ECOLOGICAL THOUGHT

LIBRI PICTURATI

A unique source for the history of plant ecology is Libri picturati A. 18–30, a collection of water-colours made mainly in the Netherlands in the second part of the sixteenth century, kept in the Graphics Department of the Jagiellonian Library in Cracow (Poland).1 More than 1,800 specimens are represented, comprising over a thousand taxa mainly from north-western Europe and the Mediterranean region, but also from Asia and America. The pictures are of such excellent quality that many of them could be used as illustrations of modern books (Figure 1).

Most of the pictures have annotations, written in different scripts, with the following information: plant names in Greek, Latin, Flemish, French, German and Italian, references to the works of ancient and Renaissance authors, and notes in Latin about localities, plant distribution, ecology, and other subjects (published in Latin2 by Ramón- Laca, 2001).

The volumes do not contain any indication of the original owners of the collection. Since many pictures match with woodcuts in various works by the famous Renaissance botanist Charles de l’Écluse (Carolus Clusius) (1526–1609), some authors have attributed them to L’Écluse (Arber, 1988: 229–231; Whitehead et alii, 1989). According to recent studies, the set of water-colours was a collaborative work by painters, plant-lovers who sponsored the paintings, and botanists (including L’Écluse) who annotated the pictures. Wille (1997), who has carefully examined L’Écluse’s life and correspondence, put forward the idea, supported by Egmont (2005), that a large proportion of the pictures were produced by order of L’Écluse’s friend, Karel van Sint Omaars (Charles de Saint Omer) (1533–1569), a wealthy plant-lover (his other name was Ranoutre – seigneur de Moerkercke, Dranoutre or Reynoutre). In his castle at Moerkercke (now in Belgium), he had a collection of curiosities and also a garden containing useful plants, both of native and foreign origin. The painters included Jacques van Corenhuyse (whose monogram can be found on three pages) and probably Pieter van der Borcht (c. 1535–1608) working for Christophe Plantin (1514–1588). L’Écluse stayed at the castle in Moerkercke, probably annotated some of the pictures, and used more than one hundred of them as illustrations for his flora of Spain and Portugal, Rariorum aliquot stirpium per Hispanias observatarum historia (L’Écluse, 1576), and a later work Rariorum plantarum historia (L’Écluse, 1601) (Ramón-Laca, 2001; Egmont, 2005). Some of the pictures were also published by Rembert Dodoens (1517–1585) in Stirpium historiae pemptades sex (Dodoens, 1583) and Matthias L’Obel in Kruydtboeck (L’Obel, 1581). According to Wille (1997), the collection was supplemented and re-arranged after 1595 by Karel van Arenberg (1550–1616), Prince and Count of Arenberg, an eminent amateur botanist, who augmented the collection with foreign plants which did not occur in the Netherlands. He also probably instructed that the collection should follow the method used by Jacques Daléchamps (1513–1588), author of Historia generalis plantarum (1586, 1587). Another hypothesis about the origin of the collection, put forward by Swan (1998), attributes the water-colours to Dirck Outgaertsz Cluyt (Theodorus Clutius) (1546–1598), but this idea is less well documented.

The water-colours in Libri picturati reflect a synthesis of art and science, so vital when they were produced (Zemanek and de Koning, 1998). Thus they are of interest to both art historians and botanists. For a long time the collection was not available to scholars. Rediscovery of the Libri picturati by Whitehead (Whitehead et alii, 1989) initiated studies of these unique pictures. Historians and art historians attempted to discover the date and place of their origin, and the names of the people associated with them (Swan, 1998; Wille, 1997; Egmont, 2005). Botanists are interested in identifying the plants depicted in this painted


“herbarium”. So far only the orchids (Künkele and Lorenz, 1990), the Umbelliferae (Apiaceae) (Baumann, 1998), and heathers (Erica spp.) from Spain (Ramón-Laca and Morales, 2000) have been examined from a taxonomic point of view.

In 2002 the Libri picturati Project was started, involving interdisciplinary studies by an international team.3 As well as research into the origins and history of the collection, and its role in the history of botanical illustrations, scholars will also investigate the plant groups, the morphology of plants, phytogeography, ecology, useful plants, and connections between the Libri picturati and l’Écluse’s garden in Leiden. It is intended to produce an annotated facsimile which will illustrate the combination of art and science, so characteristic of the Renaissance period.

ECOLOGICAL INFORMATION IN LIBRI PICTURATI A. 18–30

Pictures of plantsIn a few cases we have typical ecological pictures showing species from the same habitat on one page. However, this set of realistic water-colours, illustrating various growth forms and diverse ecological adaptations to living conditions, contains a voluminous ecological content even though contemporary authors were not aware of most of these adaptations. Almost all plants are painted at the flowering stage, with fruits quite often added, sometimes also seeds, which provide quite a rich set of phenological data.

AnnotationsFor more than half of the species ecological data were included, such as habitats, time of flowering and fruit-bearing, as well as possible cultivation in gardens, and requirements pertaining to soil, light or similar. In some cases more general observations are provided.

The most extensive written information is for species from north-western Europe, particularly Belgium and the Netherlands, and the detailed descriptions of habitats and the geographical names supplied (Frisia, Holland, and others) confirm that the plants were observed in their natural surroundings. Sometimes, the notes state “in our area” or “here it is cultivated in gardens”. In two cases, the specific dates of observations were recorded:

Cypripedium calceolus4: “This plant was seen and observed ... at the beginning of June 1564” (22/34)5

Prunus avium ‘Plena’: “In most cases it does not bear fruits, but not always, because this year, being 1564, some of the Cerasus trees had fruits” (20/79).

Arrangement of species in particular volumesTo a certain extent, ecological information is also coded by the way species are grouped in individual volumes. Some volumes, or their fragments, include smaller or larger groups of plants from certain habitats: for example, in A. 18 there are many species of the herbaceous layer of deciduous forest, as well as ruderal and meadow species: in A. 19 and 24, among others, plants of dry meadows, grasslands and shrub communities: in A. 26, aquatic and bog plants: in A. 18 and 23, synanthropic plants, those occurring in habitats created by or altered by humans. That could reflect the arrangement of the collection according to the method of Daléchamps (1586, 1587), who distinguished such ecological groups as “plants growing in marshes, ... in rough, rocky, sandy, and sunny places, ... in shady, wet, marshy, and fertile places, ... by the sea, and in the sea itself, climbing plants, spiny and prickly plants, plants with bulbs, and succulent and knotty roots” (Arber, 1988: 173). However, in most volumes


the species of various habitats are intermixed, and habitat information is contained in captions. It should be emphasised that binding of the pictures into volumes was done after the actual painting of the pictures had taken place.

PLANTS AND THE ENVIRONMENT

ClimateThere are no direct references to climate. The descriptions of habitats allow for the conclusion that the authors were aware how plants depended on light and water. These two factors were most often mentioned when habitats were described. Sites with good insolation (sunny places) were also defined as warm, whereas the shaded sites were referred to as cold or moist.

Euphrasia stricta: “in meadows and hilly sites with a lot of sunshine” (24/42).Anthericum liliago: “Occurs in elevated sites, very sunny, under thin and small shrubs” (22/24).Lavandula angustifolia ‘Alba’: “In our area, it is sown in gardens, and grows best in a very sunny place” (19/23).Asplenium trichomanes: “It likes moist and overshadowed places, growing abundantly on old fences, often on coarse walls and rocks” (18/2).

The effect of the wind was noted in one description: Milium effusum “persists for many years, but stays only in places sheltered from winds” (23/49). The information given most often is about moisture status, referred to either in general terms when a habitat is simply given as a moist or dry site – for example in the note that Phyllitis scolopendrium “favours places which abound in water” (18/4) – or in more detailed descriptions of habitats which are numerous (see below). The tendency to look for more generalised statements can also be exemplified by this note: “all Arum species grow in cold and shady places” (Arum maculatum: 22/37).

SoilUsually, two types of soil were distinguished: fertile (rich) and infertile (poor).

Centaurea centaurium: “on fertile soil ... in our area it is sown in gardens” (18/50).Isatis tinctoria, Corylus maxima: “prefers fertile soil” ( 23/107, 20/84).Senecio fluviatilis: “on mountain and fertile soil” (18/39).Prunus armeniaca: “Sown in gardens and vineyards. Prefers fertile and moist soil” (20/76).Erophila verna: “in fields and on hills, on poor soils” (18/3).

The colour or type of soil were mentioned rarely:

Vaccaria pyramidata: “prefers fertile soil, pale, muddy fields near the Mosa river, also sown in gardens” (21/90v).Ranunculus flammula: “Occurs in moist and clayey sites” (26/1).Spergula arvensis: “It occurs either spontaneously among sown cultivated plants, or is itself sown in fields, most often on dry and black soil” (18/85).

Sometimes descriptions refer to sandy sites, for example:

Veronica spicata: “Occurs in wild and sandy sites, on edges of fields and forests” (18/73).Coronopus squamatus: “along roads, on embankments, in desert and sandy sites” (18/82).Erica tetralix: “in plains and sandy hills, it also favours wooded [areas]” (20/28).

Most information pertained, however, to the level of moisture in the environment, and the description of habitats with different moisture levels were diverse (see below).


TYPES OF HABITATS AND ASSOCIATED SPECIES

Aquatic plantsMost aquatic habitats are those of inland fresh waters: ditches, slow-flowing rivers and canals. Among the pictures of aquatic plants, there are some of floating plants or those rooted to the bottom of bodies of water, for example Nymphaea alba (26/22), and others:

Nymphoides peltata: “in muddy places and in stagnant waters of Holland” (26/19).Lemna trisulca, Hydrocharis morsus-ranae, Lemna cf minor: “these three plants often occur together, floating in stagnant waters” (26/23).Hottonia palustris: “Floating in stagnant waters” (29/42).

Among the floating plants there was one carnivorous species, Utricularia vulgaris (29/40), with the following annotation: “never described before” (perhaps it is the first image of this species ever painted). Even though it was realistically represented, the trapping vesicles on submersed leaves were not shown.

Bog plantsPlants growing in shallow waters along river and canal banks, as well as in areas occupied by wetlands and bogs, are represented by a significant number of species, and the descriptions of their habitats are similarly diversified, for example:

Carex pseudocyperus: “Occurs in boggy, moist sites” (19/25v).Lycopus europaeus: “In our area it is common, occurs in boggy sites and along canals” (24/11).Veronica beccabunga, Rorippa amphibia: “on canal banks, along stagnant or slow-flowing waters” (26/87, 26/88).Ranunculus aquatilis: “It is a small herb of marshes, frequent in courtyards which are irrigated in winter” (26/29).Oenanthe fistulosa: “in wet meadows and near stagnant waters and in sites which are waterlogged in winter and dry in summer” (26/90).

For Glyceria cf fluitans (26/13) extended information was given: “Emerges in spring in shallow and boggy waters where it floats at their surface and its name derives from it, later, however, it becomes erect, forming much elongated flexible shoots”.

There is a rich set of plants occurring along banks of rivers, canals and fresh-water ditches, for example:

Iris pseudacorus: “muddy and moist sites” (22/67).Fraxinus excelsior: “Moist sites, river and canal banks” (27/18).Lythrum salicaria: “in waters, bogs, willow scrub” (26/52).Pulicaria dysenterica: “in wet areas, especially along roads, where water trickles from fields, creating a wet habitat and flows further to nearby canals” (26/40).

In one description there is a reference to the expansion of a species: Epilobium hirsutum: “Occurs naturally in very moist meadows, near ditches and streams, penetrates whenever it may grow much larger and widespread” (26/50).

Sea coast: halophytesThere are numerous pictures of halophytes, plants which occur mainly along sea coasts and thus capable of tolerating high levels of soil salinity; these pictures are probably the first of this kind. The fact that they were observed on the coasts of Holland and Zealand is confirmed in some of the captions (27/98 (Figure 2), 23/96, 26/75. In Libri picturati fifteen


Figure 2. Cakile maritima (27/98), a halophyte occurring along coasts of north-western Europe: “Occurs on the coast of Holland, near the town Sceuenynghe, with thick leaves. The whole plant is very bitter”. © Jagiellonian Library, Graphics Department, Cracow, Poland.



Figure 3. Salicornia europaea and Suaeda maritima (24/31), halophytes from coasts of north-western Europe. The annotation is one of the earliest containing information on the halophyte community: “They like very much saline soils and sea coasts, and grow in stable community, to such degree that many people considered, wrongly, there were not two, but one plant, and express such opinion”. © Jagiellonian Library, Graphics Department, Cracow, Poland.



halophytes were depicted, thirteen of them with relevant references to their habitats (Table 1). Only two species whose habitat was described as “sea coast” are not now considered as halophytes: Onobrychis sp. (O. viciaefolia or O. arenaria: 23/84) and Eriophorum angustifolium (24/8).

Three plants were depicted under two names on 24/31 (Figure 3). Under “Kali et Alkaly” there are two plants which are difficult to identify because no flowers are shown and there is no information about where they are from. If we suppose that they are from the Mediterranean area they could be identified as the diploid and tetraploid forms of Salicornia europaea. The third plant is labelled “Anthyllis Quarta”; it is Suaeda maritima. There is also a note, perhaps the earliest of this kind, about a community of sea-coast halophytes: “They like very much saline soils and sea coasts, and grow in stable community, to such degree that many people considered, wrongly, there were not two, but one plant, and express such opinion”. This annotation indicates as association similar to that formed by the halophytes Kochia hirsuta and Suaeda splendens found on the coast of Languedoc, defined during the early twentieth century by Braun-Blanquet (1931, 1933, 1947).

Table 1. Halophytes depicted in the collection Libri picturati A. 18–30.

Taxon vol./folio Habitat and locality

Anthyllis vulneraria subsp. maritima 21/89 “on sea coast and saline sites”

Aster tripolium 27/96 “at sea coast sites pounded by sea waves”

Atriplex littoralis 28/26 “on fences and in sea-coast regions. In Cilicia it

provides high yield”

Cakile maritima 27/98 “on the coast of Holland, near the town

Sceuenynghe”

Crithmum maritimum 29/38 “on sea-coast and rocky sites, sown in gardens”

Eryngium maritimum 27/51 “in wild sites and meadows, most often on sea

coasts, however”

Glaux maritima 23/96 “on sea-coast and saline areas. Known by the

residents of Zealand”

Hieracium umbellatum 28/31 “somewhere on the sea coast”

Salicornia europaea 24/31 see main text

Salsola kali 27/95 no information

Senecio crithmoides 27/96v no information

Suaeda maritima 24/31, 27/95v for 24/31 see main text; 27/95v has no

annotation

Thalictrum minus subsp. dunensis 26/75 “in sea-coast sites and on saline fields, known

in Zealand”

Thalictrum minus subsp. minus 26/76 “found on saline fields on the sea coast,

common in Zealand”

Triglochin maritima 26/14 “in sea-coast sites and very moist sites”


ForestsInformation about forests is not detailed, usually being limited to the simple term forest. Sometimes this is expanded with some additional information about the level of light or moisture, or frequency of occurrence. Examples include:

Maianthemum bifolium: “in shady forests” (18/69).Solidago virgaurea, Geranium rotundifolium, Daphne mezereum: “shady, moist forest” (18/40, 18/41, 20/38).Anemone nemorosa: “in forests and on canal banks, in overshadowed and more moist places” (26/34v).Betula cf pendula: “Occurring often in forests and in sites not subject to cultivation; prefers cold areas” (27/25).

The forest trees were only rarely named; for example, Hieracium murorum occurs in “shady forests, particularly on poor soil, often beneath an oak canopy” (28/28).

Meadows“Meadow” is noted in the annotations of many species, sometimes with a more detailed indication of the type of meadow: moist, boggy, fertile.

Cardamine pratensis: “on moist meadows and near canals and stagnant waters” (28/99).Veronica serpyllifolia: “in moist places and meadows” (18/72).Veronica chamaedrys: “mostly in meadows which are overshadowed or on slopes” (18/90).

Here are many pictures of thermophilous plants, both from dry meadows and xerothermic communities. The dry meadows are sometimes described as “wild, inhospitable places”. In the annotations, however, there are significantly fewer expressions for dry meadows than for wet ones: Ranunculus bulbosus “Prefers dry and sandy meadows” (26/32). Four members of the pea family (Fabaceae) depicted on 23/117 are correctly grouped according to the type of meadows in which they grow: “Occurs abundantly in fertile meadows” (Trifolium repens, T. pratense) and “Emerges on dried fields and meadows” (Lotus corniculatus, Trifolium dubium).

Edges of forests, meadows and fields, and hedgesThe edges of forests, meadows and fields and places “near fences” (“iuxta sepes”) (referring generally to species which form hedges or those growing at the base of fences) are set distinctly as a separate group. The abundance of this type of semi-natural habitat testifies to the remarkable level of anthropogenic transformation of the landscape at that time.

Veronica spicata: “in wild sandy places on the edges of fields and forests” (18/73).Verbascum cf lychnitis: “Occurs naturally on embankments and along the edges of fields and forests” (18/60).Rhamnus cathartica: “at fences and forest roads” (20/14).Cirsium vulgare, Carduus crispus: “along roads, fences, on the edges of fields and forests, particularly in moist and fertile places” (27/41, 27/42).

MountainsThere are many references to mountains, with most of the notes containing only general information; for example “mountains” (Polygonatum multiflorum, P. verticillatum: 30/49); “mountains, forests” (Veronica officinalis: 18/74). There are no pictures of plants occurring at high elevation; most of montane plants depicted inhabit low elevations, in the areas covered by forests, shrubs or meadows.


Figure 4. Urtica dioica (18/19), stinging nettle: “Emerges everywhere, by walls fences and in shrublands. Anyone can recognize it even in the dark only by touching”. © Jagiellonian Library, Graphics Department, Cracow, Poland.



Paris quadrifolia: “in mountains and shady forests” (18/68).Cornus mas: “In nature, they prefer mountains and hills, and they are also sown in gardens” (20/87).Juniperus communis: “Prefers mountains and hills, in our area it is rare” (27/15).

Sometimes the notes include additional remarks about light conditions, moisture levels and fertility of soil.

Castanea sativa: “Prefers mountain and shady sites” (27/4).Centaurea centaurium: “Emerges spontaneously in mountains, in well-shady and overgrown sites, on fertile soil” (18/50).Sorbus aucuparia: “Prefers mountain sites, dry and overshadowed. In our area it occurs rarely” (27/32).Cupressus sempervirens: “Prefers mountains, in our area it is an alien plant, however, it is cultivated in gardens” (27/12).Peucedanum officinale: “Emerges spontaneously in mountains; in our area it is rare, tended only in gardens” (29/27).

Occasionally the annotations describe mountain species descending into lowlands – Picea abies “likes mountains, but also descends into valleys” (27/11) – or trees which form mountain forests – Taxus baccata grows on “rocky mountains, precipices, between firs and spruces” (27/16).

Mountain ranges of various European regions, of France, Germany, Switzerland and Italy, are named: Buxus sempervirens “common in Switzerland, Burgundy” (20/19); Acer pseudoplatanus “in the higher mountains of Germany, in Belgium it occurs rather rarely on field boundary strips” (27/26); Laburnum anagyroides “common in the mountains of Italy and Narbone in southern France” (27/34); Glycyrrhiza glabra and Mandragora officinarum (see Figure 1) are “abundant in Apulia, particularly on Monte Gargano” (20/52, 30/85).

Anthropogenic habitats: synanthropic plantsNumerous notes occur about species growing in anthropogenic habitats (particularly in 18 and 23). These are indicative of the abundance in sixteenth-century Europe of plants that have for a long time accompanied human beings and which grow abundantly near human settlements. For many, the descriptions of habitats are accurate enough to designate the plants as ruderal (accompanying human settlements) or segetal (occurring as weeds in cultivated fields and gardens).

Ruderal, or nearly ruderal, habitats include the surroundings of buildings, waste lands, sites along fences and roads. Examples of ruderal species depicted include Urtica dioica (18/19) (Figure 4) and Leonurus cardiaca (18/21), as well as

Heliotropium europaeum: “elsewhere, in inhospitable places, on squares, along roads and home yards, thus in dry and sandy places; in our area it is indeed only found in gardens” (18/98).

Sisymbrium officinale: “everywhere in courtyards, archways of buildings and debris and along roads” (28/95).

In the case of segetal plants, the habitats were only referred to in general terms as “cultivated fields” or “plantations” without any precise identification of the type of plant cultivated there.

Kickxia elatine, K. spuria, Trifolium arvense: “in cultivated fields and plantations” (18/13, 18/14, 23/68).Spergula arvensis: “Emerges spontaneously in sown fields, most often on dry, black soil; much sought after as fodder for pack-animals” (18/85).Damasonium stellatum: “Occurs in Holland, but it is a nuisance to cowherds. Our cattle become fat after grazing on it but soon after suffer diseases and waste away” (26/47).


Rhinanthus sp.: “Grows in some places on high-elevated meadows and among sown cultivated plants, not without detriment to their seeds” (26/67).Apera spica-venti: “Appears among cultivated plants, not without detrimental effects thereto, and occurs often in the fields and along fences, most often in years with low yields and a lot of rain” (23/48v).

Only occasionally were the cultivated plants indicated: cereals, legumes, vegetables, and vines, for example.

Melampyrum arvense: “Grows in wheat fields, mostly on fertile soil” (23/56).Securigera securidaca: “Elsewhere emerges naturally in fields and among barley and lentil and in our area it is found only in gardens” (23/70).Veronica agrestis, V. triphyllos, V. hederifolia subsp. hederifolia, V. arvensis: “occur spontaneously among vegetables and legumes [inter olera et legumina]” (18/9: see also 18/10, Stellaria media, Arenaria serpyllifolia).Mercurialis annua: “in vineyards and gardens, among vegetables” (27/68).

For two spurges, depicted together on one sheet (27/85v), their habitat differences are correctly identified: Euphorbia helioscopia grows “in vineyards and among vegetables and legumes” whereas E. peplus occurs “for the most part around towns and on debris”. For many species the habitat was identified as boundary strips (baulks between cultivated fields), for instance for Melampyrum pratense (23/57) and Centaurea cyanus (23/67).

In gardens, besides the plants deliberately cultivated, there grew, as today, an entire array of weeds not at all desired by gardeners, as shown by this annotation for a very familiar weed: Aegopodium podagraria “occurs for the most part spontaneously in gardens and clearings in forests, where it propagates itself very widely even to a nuisance” (18/36).

Silybum marianum (27/53), a Mediterranean species cultivated in gardens, is noted as regressing to the wild state: “It occurs spontaneously in most gardens, around ruins and along fences”. Cultivation of plants and habitat requirementsFor many species, information was given about cultivation in gardens and fields. Sometimes a detailed explanation was added, such as when a plant was cultivated for medicinal, decorative or botanical research purposes. In some cases, the requirements of successful cultivation – soil type (fertile or poor), moisture level (moist or dry habitat) and light conditions (sunny or shady site) – were also given.

Fagopyrum esculentum: “Likes poor and weak soil, but can also grow in fertile soil. Sown between mid-spring and later throughout almost all of the summer; matures in the eighth or ninth week after sowing” (23/40).Setaria italica: “Likes loose soil, thoroughly tilled out but can also grow in sandy places. Sown in spring and cut – in very warm regions – after 40 days. In our area, the time for sowing is in April, while the maturing time comes in June” (23/51).Cucumis melo: “Likes sunny places, fertilized well, and soil tilled thoroughly” (28/74).

In a few cases there is information of other kinds:

Mentha aquatica: “Sown in gardens and spreads easily through roots and suckers. It dislikes manure, as well as the sun” (28/106).Ocimum basilicum, O. basilicum var. minus: “all basils are entirely summer herbs and intolerant of frost” (28/111).

The cultivation of two American plants in sixteenth-century European gardens is confirmed by such annotations as that on the portraits of Phaseolus cf vulgaris (23/87: kidney bean,


Figure 5. Phaseolus cf vulgaris (23 /87) from South America, introduced to Europe in the sixteenth century, and cultivated for its edible pod (legume): “It likes fertile soil and sunny places. In our area it is sown in gardens in April, where it can withstand ground frosts well; beans mature almost always in Autumn”. © Jagiellonian Library, Graphics Department, Cracow, Poland.



French bean) (Figure 5), and of Celosia argentea Plumosa Group (25/60: cockscomb) which was noted as liking “moist places and sticky soil (solo tenaci)”.

Errors and inaccuracies in habitat descriptionsFor a few species, the information on habitats does not agree with current knowledge. As well as the two halophytes, noted above, Helichrysum stoechas (29/45) was stated to grow on slopes and stream banks; this southern European species occurs in macchia, garrigue and in dry meadows. Umbilicus rupestris (22/45) was noted as a plant which “likes sea-side and stony places”; that is only partly true, as this species occurs spontaneously on humid shady rocks and on old walls. Aristolochia rotunda (19/61) was stated as occurring on rich soils, instead of uncultivated arid sites, and Artemisia vulgaris (19/39) as occurring on sea coasts although that habitat is not typical for that species.

Growth forms (life forms)Most of species represented in Libri picturati are herbaceous plants. Trees, shrubs and shrublets form a much smaller group (chiefly in 20 and 27), the same as creepers (several dozen species in 23, and also some in 19 and 21).

The interest in plants with beautiful flowers, which were cultivated or introduced into cultivation in the second half of the sixteenth century, resulted in an over-representation of Spring geophytes, plants growing in deciduous forests and flowering in early Spring when the absence of leaves allows the penetration of sunlight. These plants, whose shoots die after producing seeds in early Summer, persist for the rest of the year in the form of underground organs – bulbs, tubers and rhizomes. In horticulture, these organs are often used for vegetative reproduction. Volumes 21, 22 and 30 contain drawings of geophytes originating from various parts of Europe and, much less often, from Asia and America.

PhenologyPhenological data are provided in the images as well as in the captions (in about a quarter of the species). Usually, the pictures show a plant in flower, often also bearing fruits and, in many cases, also producing seeds. The intention of the authors to represent plants realistically can be seen in the manner of illustrating the shoots with fruits and seeds “cut off” from the flowering shoots (a similar way of presentation applies in modern works about plants). The annotations provide information about the time of flowering and, more rarely, about when fruits are borne or seed are mature:

Sanicula europaea: “has flowers in May and June” (18/37).Paris quadrifolia: “flowers in April, the fruit matures in May” (18/68).Agrimonia eupatoria: “begins to flower in June, the seeds mature in August” (18/23).Centaurea solstitialis: “has flowers at equinox, since its name seems to be derived from this fact” (27/54).

For some species, a note is supplied about producing flowers twice, or about dying after the period of flowering (Caltha palustris: 26/43), or after its seeds are produced (Fallopia convolvulus: 23/12).

FloweringThe volumes of the Libri appear to be an “ever-flourishing” garden. In this hortus pictus we can find plants flowering in almost every month of the year. Many of these plants are species brought from southern regions and tended in gardens. The first flowers in this “painted garden”


can be seen as early as February, and other species provide flowers without interruption until the end of the year. Several examples can be noted.

February – Lavandula multifida but only in its native place, in Spain (19/24v); Narcissus pseudonarcissus (30/48).March – there are not only bulb species blooming in March (Scilla cf bifolia: 22/8); Muscari neglectum: 30/28v) but also others such as Vinca minor (25/44).April – other bulb species such as Narcissus poeticus (30/40(39).May – numerous species, for example, Asphodelus albus (22/21).May and June – Alliaria officinalis (19/19); Sedum acre (24/17).June–July – Teucrium scorodonia (19/2); Allium vineale (22/3).Summer – in some cases the whole of the Summer is mentioned: Clinopodium vulgare (19/30). At the “end of Summer” Clematis flammula (24/47) blooms, slightly later than in eastern-central Europe (June–August) and definitively later than in southern Europe (May–July).Autumn – Scilla cf autumnalis (30/28); Colchicum autumnale: “flowering slightly before the Autumn equinox” (22/10).Winter – the palaeotropical species Withania somnifera: “in our area it flowers during winter” (30/83(12)).

Flowering of alien species acclimatized in north-western EuropeIn some cases, the author of the annotations was aware that the “local climate” can cause differences in the phenological phases: Origanum heracleoticum (19/4) “in our climate it flowers during the winter” (the steno-Mediterranean species normally flowers between June and August); maize (Zea mays cultivar: 23/38v), imported from America, “when planted in April, it ripens in August”.

Generally there is a good correlation between the phenological (mainly the flowering) periods noted in Libri picturati and the flowering period known today. Obviously, there are differences in the case of introduced species, cultivated in gardens and which have been relocated from the south to the north (Table 2). There are many examples. Salvia officinalis (19/65) is mentioned as flowering in June and July, while this steno-Mediterranean species normally flowers naturally sometime between March and May. A similar case is Umbilicus rupestris (22/45v), recorded as flowering in June and July, while this Mediterranean-Atlantic plant flowers mostly between March and June.

Table 2. Flowering period of some plants represented in Libri picturati A. 18–30 in comparison to flowering period known today in Mediterranean region.

Taxon vol./folio Flowering period Flowering period known today in Mediterranean region

Achillea ptarmica 24/43 May–September July–September

Anthericum ramosum 30/5 July May–July

Chrysanthemum segetum 25/62 June–Winter April–August

Dracunculus vulgaris 22/40 July April (–May)

Helichrysum cf italicum 29/45 “about midsummer” May–November

Hyssopus officinalis 19/32 June–July July–October

Origanum cf heracleoticum 19/4 Winter June–August

Origanum majorana 19/3 July–August June–September

Teucrium polium 19/29 July April–June

Vincetoxicum hirundinaria 24/25 June May–August


Other phenological phenomenaThere are some scattered pieces of information pertaining to other changes occurring seasonally in plants. For example, there is a note about the ferns having green leaves in Winter – “stays green throughout the year” (Asplenium trichomanes, A. ruta-muraria: 18/2; Ceterach officinarum: 18/8) – and lacking flowers – “The leaves appear in April, and die in autumn. The flowers and fruits are entirely lacking” (Dryopteris filix-mas: 18/6). There is also an interesting observation about the tufts of the pappus in representatives of the family Compositae (Asteraceae): “it has flowers in mid-Summer, the florets mature quickly and then fly away with tufts of pappus” (Erigeron sp.: 28/40). Another type of fruit, a berry, was described several times: Sambucus nigra (20/60), Viburnum opulus (20/62), Hedera helix (23/7).

Cultivated alien plants were subject to more thorough observation, and thus the annotations furnished more details: Lactuca sativa Capitata Group and Crispa Group: “Both varieties prefer well-spaced sowing in gardens with a lot of manure supplied. The plant normally flowers in the Summer, in the second or third month after sowing” (28/23).

On the sheet with Juniperus sabina (27/7v) (Figure 6) is a note discussing some

Figure 6. Juniperus sabina (27/7v) occurs on mountains in south-eastern Europe, and sometimes is planted in gardens: “With us sown in the gardens. The fruit matures towards winter time, although Guilandinus claims that the plant is sterile”. © Jagiellonian Library, Graphics Department, Cracow, Poland.



phenological matters with Melchiorre Wieland (Guilandino) (1520–1589), Professor of Padua University (Zemanek, 1998a: 104). Silybum marianum, a species described as returning to the wild state in gardens, has some more detailed notes supplied: “It flowers in June and July of the same year as it germinates, but dies shortly after producing seeds” (27/53).

Results of observations of various ecological phenomena and adaptationsAmong the plants with special adaptations, particular attention should be given to the desert xerophyte Anastatica hierochuntica (23/22v: rose-of-Jericho, resurrection-plant) (Figure 7), which dries up during drought but develops leaves immediately after watering. The annotation shows how carefully this unusual species was observed: “It opens not only during the Christmas Eve period, and not only on its own, as it was once believed by naïve local people, but always when you want it, after prior spraying with water or wine”.

Other examples of accurate observation are supplied with the illustrations of several parasitic flowering plants, including Orobanche rapum-genistae (23/94) (Figure 8), and Cuscuta epilinum (27/89) which “emerges suddenly among forbs and shrubs, particularly in flax fields and lives with their assistance, intertwined and attached to the roots of some of these plants with its projections and hairs, it overwhelms the nearest plants, saps vital stamina from them and sometimes chokes them”.

There is an interesting note to the rough hairs on the flowering spikes of wheats (Triticum spp: 23/35): “sown in Alsace and elsewhere in wooded sites – because of the roughness of

Figure 7. Anastatica hierochuntica (23 /22v), rose-of-Jericho or resurrection plant, dries up during a drought but develops leaves immediately after watering. © Jagiellonian Library, Graphics Department, Cracow, Poland.



Figure 8. Orobanche rapum-genistae (23/94) is parasitic on various leguminous shrubs in Europe: “It emerges on dry and poor soils, under a smaller shrub of Genista, which grows with considerable effort and has only some flowers or none at all. Attached to its roots via a dense network of its own roots, it drains all sap out of it and utilizes it for its own nutritional needs. It can be observed in the months of June and July”. © Jagiellonian Library, Graphics Department, Cracow, Poland.



spikes, which it grows to protect itself against grazing by wild boars”.Observations about plants germinating in various habitats are sometimes added: “They

reach variable sizes, depending on the site where they germinate” (Lotus corniculatus, Trifolium dubium: 23/117).

CONCLUSIONS

The ecological ideas contained in Libri picturati A. 18–30 provide a synthesis of the Renaissance body of knowledge about the relationships between plants and the environment. It seems that a great proportion of the data, particularly that pertaining to north-western Europe, resulted from many years of observation of plants both in the wild and in cultivation, augmented by data from the literature covering south-western Europe, in particular the Mediterranean region. The collection represents the abundance of aquatic and wetland species as well as those of moist and anthropogenic habitats including canals. The relatively high level of urban development and the disruption of the natural environment were reflected in the high number of synanthropic species. There is a striking absence of references to superstitions and magic, so often presented in the Renaissance treatises on plants (even the beliefs in magic attached to rose-of-Jericho, quoted above, are discarded as untrue by applying an experimental test). Most of the data provided in the collection are corroborated by recent knowledge, although obviously on many occasions we find the information from Libri picturati incomplete. The vast array of ecological issues contained in the collection illustrates well the process of the emergence of modern knowledge about the relationships between plants and the environ-ment. It seems that such a high level of knowledge resulted not only from the theoretical research interest of the creators of the collection, but also from the searches in the field aimed at finding new plants for introduction into gardens. An additional source of information was provided by horticultural practice which in the Netherlands of the late sixteenth century reached a very high level. It was the marriage of theory and practice, this cradle of modern science, which produced such a magnificent offspring in the form of the “painted garden” in Libri picturati.

ACKNOWLEDGEMENTS

We would like to thank Dr Jan de Koning for some unpublished data, Mr Piotr Hordyñski for acces to his unpublished manuscripts “Libri picturati A.18–30. Annotations to the pictures” and “Libri picturati A. 18–30. Technical description of the manuscripts”, and Professor Zdzisław Pietrzyk (Director of the Jagiellonian Library, Cracow) for permission to reproduce some of the Libri picturati water-colours. This publication was supported by State Committee for Scientific Research (Poland), research project No 3 P04G11424.

NOTES

1 The whole collection comprises of 16 volumes (A 16 – A 31). Since 16 and 17 contain only illustrations of animals, and 31 was illustrated later in a completely different manner, we exclude these from our considerations.

The 13 volumes of botanical pictures are bound in white vellum over pasteboard, tooled in gold. All the bindings are the same size: 51.5–52 × 36.5–37 cm.

Libri picturati A 18–30 contain 1,142 sheets, some of them with pictures on both sides (1,429 water-colours). Each painted page is separated from the next one by a blank sheet with the folio numbers written by pencil. In this paper only these numbers are included.


2 All the annotations on the watercolours in Libri picturati A. 18–30 have been re-examined by Piotr Hordyñski who has made available to us his revised (but as yet unpublished) transcriptions which correct gaps and errors in Ramón-Laca’s paper (2001). It is intended that these will be published under the auspices of the Libri Picturati Project with facsimiles of the water-colours. Our translations of the annotations are based on Hordyñski’s transcripts.

3 Heinz-Dieter Krausch (Germany); Andrea Ubrizsy Savoia (Italy); Florike Egmont, Jan de Koning, Gerda Uffelen (The Netherlands); Piotr Hordyñski, Tomasz Majewski, Alicja Zemanek, Bogdan Zemanek (Poland); Luis Ramón-Laca (Spain).

4 Species determined by Martin Christenhusz, Gerda Uffelen and Bogdan Zemanek. Plant names of European plants are according to Flora europaea (Tutin et alii, 1964–1980); of non-European plants after Bailey and Bailey (1976).

5 In this paper a shortened system of numbering is used, for example, 22/34 = volume 22 folio 34.

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Received 1 December 2005. Accepted 1 December 2006.

Documents

The beginnings of ecological thought in the Renaissance: an account based on the Libri picturati A. 18–30 collection of water-colours