Peonies of Greece

A taxonomic and historical Survey of the Genus Paeonia in Greece

William T. Stearn and Peter H. Davis

Foreword by Niki A. Goulandris

The Goulandris Natural History Museum, Kifissia- Greece 1984

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PEONIES OF GREECE

FOREWORD

Contents

Peony, queen of all herbs...

Anonymous Greek Bucolic Poet of 3^rd Century A.D.

Copyright © 1984 The Goulandris Natural History Museum, 13 Levidou Street, Kifissia, Greece Disign and format by Vasso Zambeli - Koussoula, Set up in Bodoni and Printed by “J.Makris S.A.”. Kifissias 40, Amarossion - Athens. To Angelos N. Goulandris, President of the Board of Trustees of the Goulandris Natural History Museum in Kifissia, Greece, Founder and Architectural designer of the Museum; Generous supporter of natural history studies and conservation in Greece.

FOREWORD content

My interest in the study and portrayal of our Greek peonies goes back many years to when I first had the opportunity of painting Paeonia peregrina from Macedonia and P. rhodia from Rhodes. Since then, during the past twenty years, the botanical exploration of Greece under taken by the Goulandris Natural History Museum has resulted in the amassing at Kifissia of abundant herbarium material for study and a like increase in our knowledge of the distribution of Greek plants. This has been largely due to extensive collecting by Elli Stamatiadou. Her excelent well-documented specimens of Paeonia together with the living plants introduced by her from the wild and now grown at Kifissia, made obvious the need for new taxonomic investigation of these beautiful though botanically sometimes perplexing plants.

Our long collaboration with Professor William T. Stearn, who had given much attention to Paeonia as a whole from 1935 onwards, encouraged me to ask his help in producing a detailed account of the Greek peonies to be published by the Goulandris Natural History Museum, and illustrated in colour by me. To this end Professor Stearn visited critical areas (Lefkas, Evvia, Andros and Ikaria; he had already visited Crete and Rhodes) under the auspices of the Museum to study the ecology and the variation in the wild of these plants. His experience as a botanical taxonomist, bibliographer and historian has also added much to our information regarding their long history which takes us back to remote antiquity.

On visiting the Museum some years ago Professor Peter H. Davis accepted an invitation to contribute to the volume in view of his special knowledge of the genus Paeonia in Asia Minor. He has long been concerned with taxonomic problems of the Eastern Mediterranean region, being editor and major author of the many-volume Flora of Turkey and the East Aegean Islands (1965 et seq.).

I am especially happy to have had the opportunity to work together with two outstanding British botanists on what we hope will be the definitive account of the genus Paeonia in Greece.

The Museum trusts that Peonies of Greece which has also been translated into Greek, will give the reader an opportunity to know and appreciate these regal Greek plants belonging to a genus revered over many centuries for its beauty and medicinal properties and justly called in Creek "the queen of all herbs".

Niki Goulandris Kifissia, February 1984

CONTENTS content

ILLUSTRATIONS IN COLOUR content

Plates 3-14 are reduced to 2/3 of their original size.

ILLUSTRATIONS IN BLACK AND WHITE

Figures 15, 16, 18, 21, 25-33, 35, 37 by Vasso Zambeli-Koussoula, 17 by Margaret Glaser, 19 by G.L.A. Heath, 20 by Madeleine Harley. 22, 23 by Kiki Dima and Matina Theodorou, 24 by W.T. Steam, 37 by Stella Ross-Craig.

CHAPTER I content

HISTORICAL INTRODUCTION

THE NAME PAEONIA

When the wild peonies bloom on the hills and mountains of Greece, only species of Cistus rival them in splendour of flower. A ring of golden stamens encircling crimson-tipped white-lanate carpels enhances the beauty of their broad upcurved or outspreading deep red, purplish or white petals. As long ago as the 3rd century A.D. an anonymous Greek bucolic poet extolled the paeony as "the queen of herbs" just as in China some centuries later the shrubby moutan became known as hua wang meaning "the king of flowers". Even when their conspicuous petals have fallen, peonies distinguish themselves by their elegant foliage and later by glossy black seeds resting in distinctive open crimson-lined fruits. Their roots are large and fleshy, with a somewhat unpleasant taste presumably considered indicative of medicinal virtue. Thus they must have come early to the attention of the Ancient Greek herb-gatherers, the rhizotomists or "root-cutters". Myths and superstitions collect around such plants with real or reputed curative properties; they may even have magical powers attributed to them and be associated with gods. Thus Paeonia, now a botanical generic name, is a Latin rendering of the Greek ^^^^^^^^ (paionia) and is supposed to commemorate Paeon (^^^^^^) or Paian (^^^^^^), physician of the gods and discoverer of its properties. In Greek religion and mythology the physician Paeon and the god Apollo became later assimilated into one concept, with Paian as an alternative name for Apollo, whence the word paean for a song of praise, originally a hymn to Apollo. From paeonia have come Italian, Spanish and Portuguese peonia. Old French peone and pioine. Modern French pivoine, Old English peonia and pyon and Modern English peony. William Turner (c. 1508-1568) in his The Names of Herbes (1548) stated that "Peonia is also named in greeke Glicyside, it is called in englishe peony or pyony, in duch peonienrose, in french penoisne and pinoine." There are also local variants of these, e.g. Castilian Spanish peronia and perruna, Catalan Spanish piorna, pelonia and pampalonia, German Pegunje, Pegonis, Pitonirose, Bigonje etc. (cf. Marzell, 1976) as well as numerous other names e.g. German Gichtrose (gout rose), Pfingstrose (Whitsun or Pentecost rose), French Rose de la Pentecote, Fleur de saint-Georges (cf. Rolland, 1896), in no way derived from the Greek ^^^^^^^^^^^ and Latin Paeonia.

The Greek scholar Georgios Moazzo (1893-1975) derived the names Paeon and Paeonia from the verb ^^^^^^^^ (I strike). As part of the psychological treatment which obviously was important even in ancient medicine, the medical practitioner or shaman struck his patient with a magical wand while uttering the incantation ^^^^^^^^^^ (o you Paeon) which became associated with healing and with Apollo. Partridge (1966) gives the same derivation.

The mongrel English spelling paeony mixing the centuries-old traditional English peony and the Latin paeonia was a 19th-century introduction, possibly first used by G. Anderson, with nothing to commend it, and found no acceptance in America.

MYTHOLOGY content

Plants of wide distribution and distinctive appearance acquire a diversity of local names; these may persist in local speech or be copied from one book to another long after one of them such as paeonia has attained standard use. In discussing far-fetched notions Theophrastus (c. 370-288 B.C.) instanced a superstition concerning ^^^^^^^^ which some people then called ^^^^^^^^^^^^ (glucuside): this plant "should be dug up at night, for if a man does it in the day-time and is observed by woodpecker ... he risks the loss of his eyesight" (^^^^^^^^^^^^^^^^^^^^ ^^^^^^^, Enquiry into Plants IX, cap. 6, transl. Hort). Pliny, who in the 1st cent. A.D. rendered so much Greek lore into Latin, repeated this: "praecipiunt eruere noctu, quoniam si picus Martios vident tuendo in oculos impetum faciant'¹"' (Nat. Hist. X-XIV, cap. 10). Pliny's Elizabethan translator Philemon Holland rendered that passage in 1601 as "It is said, that this hearbe must be gathered in the night season; for if the Rainbird, Woodpeck or Hickway, called Picus Martius, should chance to see it gathered, he would flie in the face, and be readie to peck out the eyes of him or her that had it." This appears to be the only record of the woodpecker as a defender and conservationist of peonies. Such beliefs, like those relating to the mandrake, must have been spread and perpetuated with a practical motive by the ancient herb-gatherers who knew them to be false; they guarded medicinally and commercially valuable plants against undue exploitation by the uninitiated and also maintained a trade monopoly. Accordingly they testify to the former importance of the plants concerned, notably paeonia and mandragora. Continuously copied, they survived in literature, though not necessarily believed, to the 16th century. The English barber-surgeon, herbalist and gardener John Gerard (1545-1612) dismissed them contemptuously in his Herball (1597) with the remark: "The like fabulous tale hath been set forth of mandrake. But all these things be most vaine and frivolous, for the roote ofpeonie, as also the mandrake, may be removed at any time of the yeare, day or houre whatsoever".

Greek legend, preserved in works attributed, probably spuriously, to Hesiod and Theocritus but beneath the notice of Theophrastus and Dioscorides, invented a rivalry between Paeon who knew all the remedies and his master and former teacher Aesculapius, who, jealous of Paeon, occasioned the latter's death. Paeon had, however, cured Hades (Pluto) of a wound inflicted by an arrow-shot from Hercules and Hades transformed him into a herb thereafter named ^^^^^^^^. According to another legend, Leto, the mother of Apollo, revealed to Paeon the virtues of a herb on Mount Olympus for easing childbirth and this was later named ^^^^^^^^^^^^ in his honour.

DIOSCORIDES' ACCOUNT OF PAEONIA content

The earliest surviving account of peonies is to be found in the Greek herbal of Pedanios Dioscorides (1st cent. A.D.), ^^^^^^^^ ^^^^ ^^ called in Latin De Materia medica. Here (III, cap. 157) two kinds are distinguished by their foliage, the "male peony" (^^^^, paionia arren) and the "female peony" (^^^^, paionia theleia): the male peony had leaves like walnut, i.e. rather coarse, imparipinnate with distinct broad leaflets, and the female had leaves like smyrnium, i.e. much more divided. This use of the terms "male" and "female", perpetuated in such modern botanical plant-names as Dryopteris filix-mas, Paeonia mascula, Cornus mas and Anagallis foemina, was purely metaphoical without any implication of actual sexuality in plants (cf. Saint-Lager, 1884). It is of great antiquity. The most obvious example of two kinds resembling each other closely but nevertheless distinct is provided by the separation of the sexes in mankind and domestic animals. Hence to distinguish nomenclaturally the members of a pair of species of plants, the more robust or coarser kind or with hard wood was designated as "male", the more slender or delicate kind or with softwood as "female". Likewise, as in Anagallis, the red-flowered plant was designated "male" and the blue-flowered plant "female", or, as in Symphytum, the purple-flowered "male", the white "female".

The text of Dioscorides (see Appendix) was copied and copied throughout the Middle Ages down to the invention of printing in the 15th century in both Greek and Latin versions, inevitably differing here and

there and diverging from the original by the occasional addition or omission or by the re-arrangement of material. Painstaking comparison of numerous manuscripts enabled Max Wellmann to establish what scholars now accept as the authentic Dioscoridean text. To this were added, at an early period, synonyms derived from another source, a now lost work attributed to Dioscorides' contemporary Pamphilos, and also illustrations derived from several sources, which are often valuable for indicating the traditional applications of Dioscoridean names. One illustrated manuscript of special importance is the Codex Aniciae Iulianae nunc Vindobonensis made in Constantinople in the 6th cent. A.D. Some of the illustrations in this Vienna Codex were lost during its eventful history (cf. Stearn, 1976), among them being those of Paionia. These illustrations have been preserved in some other Dioscoridean codices, notably the Codex Patavinus of 1350 in the Seminario Vescovile of Padua (cf. Stannard, 1971:178) and the 15th century Codex Chigianus in the Vatican Library, Rome. On the page (fol. 193 verso) relating to paionia arren and paionia theleia in the Padua codex 194 there are two illustrations (Frontisp.). The upper one has slender roots, as described by the herbalist for the "male peony", and the lower one has swollen roots, as described for the "female peony". Although the drawings of the leaves have been much simplified and conventionalized, the two or three spreading carpels definitely establish them as the earliest known recognizable portrayals of peonies. Regarding this codex, cf. Mioni (1959), Pächt (1975).

The distinction in leaf made by Dioscorides between the male peony (paionia arren) and the female peony (paionia theleia) corresponds essentially to that made in F.C. Stern's Study of the Genus Paeonia (1946) between subsect. Foliolatae (typified by P. mascula) and subsect. Dissectifoliae (typified by P. officinalis sensu stricto). Dioscorides as a practical herbalist contented himself with the medicinal use of peonies. An addition to his work stated that the peony should be dug up before the rising of the sun and is good against poisons, witchcraft, fevers and the assaults of devils. Its mythology became entangled with that of the mandrake and an elaborate ritual became associated with its uprooting. A Greek poem, quoted by Anna J. Papamichael (1975), even stated that "the peony is the queen of all herbs".

MEDICINAL REPUTE content

The former medicinal repute of peonies, as of mandrake (cf. Stearn, 1976), cannot be dismissed as groundless even though a reading of old herbals notably, the medieval Livre des Simples Medecines (cf. Opsomer etc., 1984), gives the impression that almost every plant was effective against almost every ailment in some way or other and so leads to the opinion that it probably cured none. Certainly the former high repute of such a plant as betony, Herba Vettonica (Stachys officinalis), seems to us most astonishing. Nevertheless among so many plants now known to be medically useless were some which remain important and a few which, though, unlike them, are no longer included in official national pharmacopoeias, have properties which made them valuable. Thus the roots of peonies (pharmaceutical Radix Paeonine) contain a substance at one time considered an alkaloid and named ‘peregrinin’ and ‘paeonin’, apparently a glycoside, which yields an oil (Paeonol), and have a limited use even now, as have the flowers (Flores Paeoniae) and seeds (Semen Paeoniae). The characteristics of the roots are described in detail in H. Thoms, Handbuch 5: 819-821 (1929) and with less detail in Potter's New Cyclopaedia of Botanical Drugs and Preparations, 8th ed., 323 (1968) by R.C. Wren: "the root appears in commerce in scraped, spindle-shaped pieces, averaging 3 in. long and ½ - 2/3 in. in diameter, pinkish grey or dirty white, strongly furrowed, and shrunken longitudinally. The transverse section is starchy, radiate with the medullary rays more or less tinged with purple." It retains its ancient reputation for good antispasmodic and sedative properties, according to P. Fournier, Le Livre des Plantes medicinales 3: 237-239 (1948) and R.C. Wren, loc. cit. (1968). The later states that peony root has been “successfully employed in convulsive and spasmodic nervous affections such as chorea, epilepsy, spasms etc.” taken as an infusion of the powdered root in boiling water. Thus the ancient association of peonies with Paeon the healer has a better medicinal foundation than have many herbs known from antiquity through the work of Dioscorides and his medieval successors. Apparently, however, their use lias disappeared from modern Greek folk medicine.

PLINY'S ACCOUNT OF PAEONIA content

The Roman encyclopedist Pliny the Elder (1st cent. A.D.) was a contemporary of Dioscorides but, despite 106 almost identical passages in their works listed by Max Wellmann (1889), there is no reason to suppose that either author knew of the other: both drew upon the same earlier Greek sources, notably a now lost work of the 1st century B.C. by Sextius Niger, according to Wellmann. Pliny too included paeonia and, like Dioscorides, gave pentorobon and glycyside as alternative names. His account, as translated by H.S. Jones (1955), reads: "The first plant to be discovered was the peony which still retains the name of the discoverer [Paeon]; it is called by some pentorobon, by others glycyside, for an added difficulty in botany is the variety of names given to the same plant in different districts. It grows on shaded mountains, having a stem among the leaves, which bears on its content four or five growths like almonds, in them being a large amount of seed, red and black. This plant also prevents the mocking delusions that the Fauns bring on us in our sleep. They recommend us to uproot it at night-time, because the woodpecker of Mars, should he see the act, will attack the eyes in its defence" (Nat. Hist., XXV, cap. 10; Loeb Class, Lib. 7: 157; 1955).

THE 15TH CENTURY content

During the Middle Ages knowledge of peonies did not advance beyond the statements of Dioscorides and Pliny, but two species native to Southern Europe and now named Paeonia officinalis L. sensu stricto (P. officinalis var. alpha feminea L.) and P. mascula (L.) Miller (P. officinalis var. P mascula L., P. corallina Retzius) were known to medieval herbalists and one (P. officinalis, sensu stricto) became commonly grown in Central European gardens, presumably more because of its decorative red flowers and foliage than for attributed medicinal properties. Both came to the attention of the 16th-century herbalists who have been called the "fathers of botany" and also of artists. Thus Albrecht Dürer's delicate watercolour, "Virgin and Child with a Multitude of Animals" ("Maria mit vielen Tieren") painted about 1503, now in the Albertina, Vienna, portrays a peony, undoubtedly P. officinalis (sensu stricto). The first printed works to mention Paeonia were simply Pliny's Historia Naturalis (1469) and d'Abano's translation into Latin of Dioscorides' Materia Medica. (1478). These were followed by the Herbarium Apulei Platonici (1481) published in Rome by J.P. de Lignamine (C.F. La Legname) from a medieval manuscript, apparently of the 9th century A.D., found in the library of the Montecassino abbey. The relevant entry begins: “Nomen Herbae Peoniae. A graecis dicit Pentorobon. Allii Cudionena. Alii Agbaosotes. Itali Peoniam. Iventa Peonio nomen auc-toris retinet. Nascitur crete”. The accompanying crude illustration (Fig. 2) bears no resemblance whatever to any peony; indeed virtually all the figures in this work are useless as guides to the plants concerned, the result of continual mindless copying over a long period of time.

This is also exemplified by the fanciful illustration (Fig. 3) to text on Paeonia in Tractatus de Virtutibus Herbarum (1499) reproduced in Herbolario volgare (1522).

Not long after the publication of the Herbarium Apulei Platonici, the printer Peter Schöffer published in Mainz in 1484 the first German-printed herbal which, having no title, is variously known as Herbarius, Herbarius Moguntinus, Aggregator practicus de Simplicibus and Latin Herbarius with Latin text and woodcut illustrations supposedly derived from a 14th-century manuscript source. It contains no illustration of a peony. In 1485 Schöffer published in Mainz a much bigger differently illustrated herbal with German text variously known as Der Gart, Gart der Gesundheit, Herbarius zu Teutsch and German Herbarius. Illustrations were evidently drawn from nature specially for this work by Erhard Rewich and they were copied in others; as stated by Agnes Arber (Herbals, 2nd ed. 26; 1938), “no work which excelled or even equalled them was produced, until a new period of botanical illustration began with the herbal of Brunfels, published in 1530.” These include one of a peony clearly recognizable as Paeonia officinalis (Fig. 4). The degeneration which such an illustration could suffer through unskilful copying is exemplified by the crude version of the same woodcut (Fig. 4) in the 1497 Strasbourg edition by Johann Prüs of the Ortus Sanitatis first published in 1491 by Jacob Meydenbach. (cf. Anderson, F. 1983).

THE 16th CENTURY content

The first stage in the development of modern botany was thus the publication in print of the works of Pliny, Dioscorides and Theophrastus which had hitherto only been available here and there in laboriously copied manuscripts. This was almost entirely the work of Italian printers, editors and commentators. "The second stage in the post-medieval evolution of taxonomic botany", as stated by Sprague and Nelmes (1931), "began when it was realized that much of the earlier work was relatively valueless, owing to doubts regarding the identity of the plants concerned. Here German botanists took the lead. The need for precision was met by the provision of recognizable illustrations in the herbals of Brunfels and Fuclis." Otho Brunfels (1488-1534) provided no illustration of a peony, but Leonhart Fuchs (1501-1566) did. Fuchs, who from 1535 to 1566 was professor of medicine at the University of Tübingen, published his De Historia Stirpium in Latin in 1542 and a German edition, New Kreuterbuch in 1543, both of which provide the same woodcut illustration of a peony (Hist. 202; Kreuterb. t. cxii), coloured in some copies, under the names Paeonia femina and "'Peonienbluom, Gichtwurz". He stated that it was planted in all gardens in Germany. It is the same species with dissected leaves portrayed by Dürer. Linnaeus specially quoted this illustration (Fig. 5; Plate 1) as representing his Paeonia officinalis (alpha) feminea and thus unambiguously establislied the application of the specific name.

Fuchs's Italian contemporary Pierandrea Mattioli (1500-1577) published in Czech in 1562 a herbal, Herbari Ginak Bylinarz, with an elaborate woodcut of the same species under the name Paeonia. Mattioli used the same illustration (Fig. 7) in his Commentarii in sex Libros Pedacii Dioscoridis (1565), relabelling it Paeonia foemina, and added another woodcut (Fig. 6) under the name Paeonia mas which portrays the species now called P. mascula (P. corallina). Both species have purplish red flowers. Mattioli’s illustrations exhibit very elaborate shading, whereas the earlier woodcuts of Fuchs consist of plain outlines without shading and hence could be coloured. Mattioli recorded P. foemina as common in Italy but P. mas as found in a few places. For economy the artists of Fuchs and Mattioli portrayed plants in flower and fruit at the same time. Although all the European peonies bear only one flower to a stem, Mattioli’s artist improved on nature by making their stems both several-flowered and branched.

Excellent little woodcuts (Figs 8, 9) illustrating Paeonia officinalis, including the old red double form 'Rubra', and P. mascula were published in the Plantarum seu Stirpium Icones (1581) of Matthias de l’Obel (Lobelius, 1538-1616) and make clear, as did the earlier woodcuts of Mattioli, the difference in roots between P. officinalis, which has them markedly swollen, and P. mascula, in which they are tapering, more slender and carrot-like. Linnaeus chose the figure on p. 684 right (Paeonia mas) as a good representation of his P. officinalis (beta) mascula. L'Obel had earlier used these woodcuts (with the exception of P. mas cum semine, p. 685) in his Plantarum seu Stirpium Historia 389-391 (1576) and also (with P. mas cum semine included) in his Dutch edition Kruydtboek 830-832 (1581).

Although Dioscorides was presumably acquainted with peonies in Greece and Asia Minor, the first definite record of a peony on Greek territory stands to the credit of Pierre Belon (1517-1564), a French naturalist who travelled in the Near East between 1546 and 1549 and survived all the many hazards of his journeys only to be murdered in Paris. In 1553 he published Les Observations de plusieurs Singularitez et Choses mémorables trouvées en Grèce, Asie, Judée, Egypte, Arabie et autres Pays estranges, a highly interesting travel-book (Fig. 10). He had visited Crete and recorded that here grew a white-flowered peony, undoubtedly the species later named P. clusii (Fig. 11). The island of Crete was then under Venetian rule. The next reference to a white peony occurs in a letter, printed in 1601, from a Venetian doctor, Honorius Bellus (Onorio Belli; d. 1604) of Vicenza who lived at Chania (Cydonia) in Crete from 1583 to 1596. Bellus corresponded with the great Flemish botanist Carolus Clusius (Charles de l’Escluse, 1526-1609) who appended these letters to his Rariorum Plantarum Historia (1601). In a letter of 23 September 1593 Bellus informed him that a white-flowered peony grew commonly in valleys of the highest mountains of Crete and that he had seen neither the male nor the female red-flowered peonies on the island. Nothing more was recorded about this Cretan peony (P. clusii) until the 19th century. Clusius accordingly added this information under the name Paeonia Cretica to the account of Paeonia in his Rariorum Plantarum Historia 1:281 (1601). This account begins with a note on Paeonia byzantino semine nata having a flower with eight, ten or more red (not purple) petals, accompanied by a woodcut (Fig. 12), which is the first illustration of the Balkan species later named P. peregrina (P. decora). Clusius had, however, described it earlier in his Rariorum aliquot Stirpium per Pannoniam 401 (1583). In 1601 he also described eight other kinds of Paeonia, including one from Spain (P. officinalis subsp. humilis) and two double-flowered forms.

THE 17th AND 18th CENTURIES content

By 1623 when Caspar Bauhin published his [Pinax] Theatri Botanici listing about 6000 kinds of plants with their synonyms, a work it had taken him 40 years to complete, ten kinds of single peonies and four double forms had become known. These included

I. Paeonia folio nigricante splendido, quae mas (now P. mascula subsp. mascula);

II.Paeonia communis vel foemina (now P. officinalis subsp. officinalis);

III. Paeonia tenuis laciniata subtus pubescens flore purpureo (now P. officinalis subsp. humilis.)

IV. Paeonia folio subtus incano, flore albo vel pallido (which included P. clusii) and

V. Paeonia peregrina flore sature rubente (now P. peregrina). Several of these peonies were illustrated by large copper engravings in Besler's Hortus Eystettensis, Vern. Ordo 6 (1613) drawn from plants cultivated in the magnificent garden of the Prince Bishop of Eichstatt, Johann Konrad von Gemmingen. The preparation of this work, according to Besler himself, took 16 years; it contains 374 plates representing 667 species. The peonies illustrated include the Balkan species from Constantinople (earlier Byzantium, now Istanbul), P. peregrina (Fig. 13).

The Paradisi in Sole Paradisus terrestris. A Garden of pleasant Flowers (1629) by John Parkinson (1567-1650) was long the best known English gardening book and it retains its interest as a delightful first-hand record of plants cultivated in England early in the 17th century. Parkinson's account of Paeonia (pp. 341-344) begins with the statement: “There are two principall kindes of Peonie, that is to say, the Male and the Female. Of the Male kinde, I have onely known one sort, but of the Female a great many: which are thus to be distinguislied. The Male his leafe is whole, without any particular division, notch or dent on the edge, and his rootes long and round, divided into many branches, somewhat like to the rootes of Gentian or Elecampane, and not tuberous at all. The Female of all sorts hath the leaves divided or cut in on the edges, more or lesse, and hath alwaies tuberous rootes, that is, like clogs or Asphodill rootes, with many great round peeces hanging, or growing at the end of smaller strings, and all ioyned to the contentpe of the maine roote”. Parkinson then described six kinds:

1. Paeonia mas. The Male Peonie (now Paeonia mascula);

2. Paeonia femina vulgaris flore simplici. The ordinary single female peonie (now P. officinalis);

3. Paeonia femina vulgaris flore pleno rubro. The double red peonie;

4. Paeonia femina flore carneo simplici. The single blush peonie;

5. Paeonia femina flore pleno albicante. The double blush peony;

6. Paeonia femina Byzantina. The single red peony of Constantinople (now P. peregrina).

He then stated that "All these peonies have beene sent or brought from divers parts beyond the seas; they are endenized in our Gardens, where wee cherish them for the beauty and delight of their goodly flowers, as well as for their physical vertues”.

Later works down to the 18th century added no information relevant to the peonies of the Balkan Peninsula. An engraving (Fig. 14) of Paeonia hyemalis pumila, Rosae rubrae in some copies of Francesco Cupani’s posthumus and extremely rare Panphyton Siculum (1713) represents the dwarf Sicilian peony (now P. mascula subsp. russi) named P. russi by Bivona-Bernardi in 1816. This central Mediterranean insular species occurs in western Greece (Fig. 23).

For Carl Linnaeus (1707-1778) all the kinds of peony distinguished by his predecessors belonged to one species which he named Paeonia officinalis in his Species Plantarum 1:530 (1753), the starting point of modern botanical literature: “limites inter species non reperi, hinc conjunxi” he stated, “I have not discovered limits between species, hence I have united them”. His specific epithet officinalis refers to their medicinal repute. He maintained, however, the traditional distinction between the female and male peonies derived From Dioscorides by treating them as varieties, one being named var. (alpha) feminea the other var. (beta) mascula. Later authors separating them as species have restricted the specific name P. officinalis to his var. (alpha) the plant illustrated in 1542 by Fuchs, whose woodcut he selected as an outstanding representation.

Linnaeus’s older contemporary Philip Miller (1691-1771), from 1722 to 1770 curator of the Chelsea Physic Garden, possessed an almost unrivalled knowledge of plants in cultivation, much of which he incorporated in successive editions of his Gardeners Dictionary, from 1731 to 1768. Well versed in the classifications of John Ray and Joseph Pitton de Tournefort, he accepted reluctantly the classification and binomial nomenclature of Linnaeus and rejected many of Linnaeus's broad generic and specific concepts which conflicted with his own firsthand observation and judgement. He recorded twelve kinds of Paeonia, both single and double, in the 6th edition (1752) of his Dictionary. He ignored Linnaeus's view that such plants differing between themselves in root and leaf were variants of one species. In the 7th edition (1759), which provides an amplified account of the genus, he referred them to six species distinguished by diagnostic phrase-names. He maintained the same six species in the 8th edition (1768), published when he was 77, but, having at last accepted Linnaean binomial nomenclature, he added specific epithets, namely 1 (Mascula), 2 (Foemina), 3 (Peregrina), 4 (Hirsuta), 5 (Tatarica), 6 (Lusitanica). Thus, tardily. Miller became the author of the accepted names P. mascula for the “male peony” later named P. corallina by Retzius, and P. peregrina for the Balkan peony of Clusius, later named P. decora by George Anderson.

THE 19th AND 20th CENTURIES content

Meanwhile the number of kinds of Paeonia in gardens so increased that early in the 19th century Joseph Sabine assembled 70 kinds under different names in his garden at South Mimms, Barnet, near London. He and his friend George Anderson allocated them to 13 species, which the latter described in “A monograph of the genus Paeonia” published in Transactions of thie Linnaean Society of London 12: 248-283 (February 1818). Anderson, obviously both a keen and competent gardener and botanist esteemed by his contemporaries, was killed by a fall from his carriage on 10 January 1817, two days after completing his manuscript. To this Sabine added a postscript (pp. 283-290) dated “20 December 1817” correlating Anderson’s work with the account of Paeonia published meanwhile by A.P. de Candolle in his Regni Vegetabilis Systema naturalis 1: 386-394 (November 1817). Anderson gave detailed descriptions, all based on garden plants, and extensive synonyms and thereby made a notable contribution to the understanding of the genus, but his statements on distribution were taken from the literature, some of it erroneous. He described Clusius’s red peony from Constantinople (P. peregrina) as a new species, P. decora, appropriately named indeed, and another, of unknown geographical origin, P. arietina, the large recurving follicles suggesting a ram's horns.

Botanical exploration of the Balkan Peninsula had hardly then begun. Sibthorp and Smith's Prodromus 1: 369-370 (1809) recorded P. officinalis from Crete, this undoubtedly referring to P. clusii, and P. corallina from Zante (Zacynthos), presumably P. mascula subsp. russi. Even Boissier in his Flora Orientalis 1:97 (1867) had little more to add; he named the Cretan peony (P. clusii) as P. peregrina var. glabra and one from Mount Parnassus (P. parnassica) as var. latifolia and recorded P. decora (i.e. P. peregrina) from “Rumelia”, the central Balkan region.

The truth of Boissier’s remark “species inter se valde affines difficile delimitandae” (the species closely akin to one another difficult to delimit) has been evident to every student of the genus from Linnaeus onwards.

Franz Wilhelm Sieber (1785-1844) from Prague, who made a collecting journey to Crete in 1817, gathered then the white Cretan peony and from his material his compatriot I.F. Tausch (1793-1848) named it P. cretica in 1828. Unfortunately Sabine had validly published in 1824 the name P. cretica for a pink-flowered peony grown in the Oxford Botanic Garden which was not of Cretan origin. Since the Cretan peony lacked a legitimate name I (W.T.S.) accordingly named it P. clusii in honour of Charles de l’Escluse and this name with which I had annotated specimens in many herbaria from 1935 onwards was validly published by F.C. Stern in 1940 and K.H. Rechinger in 1943.

Anderson’s successor as monographer of the genus Paeonia in gardens was Richard Irwin Lynch (1850-1924), a Kew-trained gardener from 1879 to 1919 Curator of the Cambridge University Botanic Garden. He immensely enriched the garden's stock of plants and became particularly interested in peonies. His study of these in a living state enabled him to publish “A new classification of the genus Paeonia” in Journal of the Royal Horticultural Society of London 12: 428-445 (November 1890). This was a valuable synopsis for horticultural use but added nothing to knowledge of the Balkan peonies, of which the only one then introduced was P. peregrina, described by Lynch as P. officinalis var. lobata and attributed to Portugal.

The work of Anderson and Sabine having been based on horticultural material (as was that of Lynch), a German high school teacher Ernst Huth (1845-1897) published a “Monographie der Gattung Paeonia” in Engler's Botanische Jahrbücher 14: 258-276 (1891), giving special attention to distribution testified by herbarium material at Berlin. Horticulturists familiar with living plants, which in gardens may only be individuals of one clone or seedlings from one introduction only, with restricted variability, accordingly tend to emphasize differences of habit and flower colour, which may not be evident in dried specimens, and they often adopt narrow specific concepts; herbarium botanists on the other hand tend to adopt much broader specific concepts, sometimes excessively broad, through lack of observable differences in dried specimens and through acquaintance with representatives of more wild populations than occur in gardens.

Huth distinguished 15 species, some being very broadly defined. Thus within P. corallina he included var. (alpha) typica (now P. mascula subsp. mascula); (beta) flavescens (Presl) Huth (now P. mascula subsp. hellenica); (gamma) pallasii (now P. mascula subsp. triternata); (delta) broteri (Boiss. & Router) Huth (now P. broteroi); (epsilon) russi (Bivona) Huth (now P. mascula subsp. russi); (theta) cambessedesii (Willk.) Huth (now P. cambessedesil). He maintained P. decora (now P. peregrina, with P. romanica as a synonym) as an independent species and may have been the first to record it precisely in a truly wild state from Serbia. His P. peregrina included var. (alpha) officinalis, (beta) villosa (P. paradoxa DC); (gamma) humilis (Retzius) Huth (now P. officinalis subsp. humilis); (delta) cretica (now P. clusii); (epsilon) banatica (Rochel) Huth (now P. officinalis subsp. banatica). Such extreme “lumping” has had few followers but J. Cullen and V.H. Heywood have come near to it in their “Notes on the European species of Paeonia” in Feddes Repertorium 69: 32-35 (1964).

Down to 1899 the name P. peregrina had been applied to peonies of south-western Europe now referred to P. officinalis (including humilis). When investigating Balkan peonies Carl Fritsch concluded that the name P. peregrina belonged rightly to the Balkan species from Constantinople named P. decora by Anderson in 1818 and P. romanica by Brandza in 1879 and he published a note on this in Verhandlungen Zool.- Bot. Gesellschaft Wien 49: 240 (1899). Acceptance of the name in this sense by Otto Stapf in Curtis's Botanical Magazine 144: t. 8742 (1918) has led to current usage. Fritsch rejected the Linnaean name P. officinalis and used the name P.feminea (L.) Desf. for the female peony (P. officinalis, sensu stricto) and P. mascula (L.) Desf, for the male peony, with P. corallina as a synonym.

In 1903 Camille A. Jordan published the third and last volume of Icones ad Floram Europae novo Fundamento instaurandum spectantes by Alexis Jordan (1814-1897) and Jules Fourreau (1844-1870), the publication of which had been interrupted by the death of Fourreau in the Franco-German War. This volume contains descriptions by Alexis Jordan and coloured plates by C. Delonne of six peonies native to France and Corsica. Jordan was a strongly religious botanist and horticulturist with a keen observant eve for minute differences among closely allied plants and hence an extremely narrow species concept, for he disbelieved in evolution and, on the contrary, believed that these microspecies or forms had been so created by God and had ever after retained their characters unchanged. Their distinction necessitated the publication of scrupulously accurate and detailed illustrations. The six beautiful plates of Paeonia in Jordan and Fourreau’s Icones (3: 37-38, tt. 318-323; 1903) rank botanically as the best ever published for their wealth of detail. Jordan's P. glabrescens and P. revelieri both from Corsica are now included in P. mascula subsp. russi; his P. leiocarpa and P. modesta both from Pyrenees Orientales in P. officinalis subsp. humilis; his P. monticola and P. villarsii both from Hautes-Alpes in P. officinalis subsp. officinalis. The value of this work as regards Paeonia is accordingly in the high quality of its illustrations. Paeonia mascula subsp. russi has its most eastern stations in western Greece; Jordan's illustrations portray it from the western extreme of its range (plate 2).

Otto Stapf (1857-1933) greatly influenced later work on the genus Paeonia, which he studied critically at Kew for many years, although he never lived to complete an intended revision. He was born in Ischl, Austria, studied at Vienna under Anton Kerner von Marilaun, whose research assistant he was from 1882 to 1889 and whose narrow specific concepts he imbibed, and then in 1891 came to England for employment as an assistant in the Herbarium of the Royal Botanic Gardens, Kew. From 1909 to 1922 he was Keeper of the Kew Herbarium. In his retirement he edited Curtis's Botanical Magazine vols. 148-156 (1922-1933), for which he wrote most of the text, including that on Paeonia tomentosa in vol. 155: t. 9249 (1931). His careful identifications and annotations of the specimens in the Kew Herbarium and his discussion under t. 9249 of the taxonomic importance of the carpels bear witness to his intimate understanding of the genus. This discussion merits reprinting here:

“A glance at the present plate and at t. 6645 of the Botanical Magazine will reveal a striking resemblance between the plants figured, the only obvious difference being the presence of a dense tomentum in the carpels of the former and its complete absence in the latter. Other minor differences there are, as in the colours of the stamens and leaves and their venation, but they are slight. The contrast in the carpels, however, is important and it constitutes a problem which recurs in connection with other peonies and affects their taxonomic status and na ming. There are, in Paeonia, at least four pairs of types whose members are stated to differ almost solely in the presence or absence of a tomentum to the carpels. I enumerate them here adding the areas they inhabit:

“The contrast between these types is the more striking as no intermediate states seem to occur. They are mutually exclusive over large areas and may therefore be considered as genetically fixed within these areas. At the same time they are coexistent in other portions of their respective areas; but whether they occur there in close proximity and in what proportion, or whether they inhabit edaphically identical stations we do not know. From this limited geographical association some authors have concluded that each such pair represents one ‘variable’ species. Now all the plants enumerated above have been in cultivation, and most of them for a long time, without showing — so far as our records go — any abrupt or gradual change from one type to the other, so that we may assume that the differentiation into contrasting types is under all circumstances highly stable. At the same time it might be argued, that, although these eight peonies represent definite fixed types, those of each pair have a common and recent origin. As, however, a sharp distinction of species with glabrous and with tomentose carpels runs through the remainder of the genus though not resulting in the appearance of parallel forms tike those under discussion, we may assume that the causes, which underlie this phenomenon of parallelism, date far back in the history of the genus.

“Space forbids to discuss the problem referred to in detail in this place. It may suffice to state that a careful examination of those plants in the herbarium and in the living state has convinced me that they are not all ‘parallel’ to the same degree, and that they ought to be accepted as specifically distinct. The parallelism is least pronounced in the P. Russi - P. Corsica, so that it is surprising that its components should ever have been linked as subspecies or varieties of one species, whilst it is undoubtedly manifest in the other three pairs.”

Stapf’s association with the Royal Horticultural Society and his unique standing as a botanical authority on Paeonia led Frederick Claude Stern (1884-1967), a wealthy merchant banker and former army officer with a celebrated garden, to propose that they should produce a joint work, Stapf to contribute a taxonomic revision, Stern the horticultural notes summarizing his experience in growing peonies from 1919 onwards in his chalk garden at Highdown, near Worthing, Sussex. England. Stern had found that they throve there and he enthusiastically acquired as many species as possible for cultivation and study. His article on “Paeony species” in Journal of the Royal Horticultural Society of London 56: 71-77 (1931) begins: "I intend to talk about Paeony species and I intend to talk about them from a gardener's point of view. I am not a botanist and therefore I can only describe them as a gardener and say how they grow in my own garden. For many years past I have raised all the Paeony species I could get hold of from seed; but I find the naming of them, especially some of the European ones, exceedingly difficult, and I also find that the botanists differ as to the correct naming of some of them ... Perhaps this discussion will encourage the botanist to produce a scientific monograph on this genus which is up to date and easy for the amateur gardener to understand." He employed the celebrated botanical artist Lilian Snelling (1879-1972) to paint them from life for this book. Collaboration between Stapf the scholar and Stern the gardener would have been ideal. Stapfs sudden death on 3 August 1933 at Innsbruck while on holiday there temporarily ruined Stern's plan. Having Lilian Snelling's beautiful paintings on hand he approached me to prepare the botanical text which would enable them to be published, and I agreed. Taking Stapfs herbarium determinations as a basis I worked out the synonymy of the taxa he had recognized and studied the herbarium material at Kew, the British Museum, Edinburgh, Prague, Vienna and Goteborg as well as many living plants. I had assumed Stern wanted me to do the work that Stapf would have done. I found indeed that he wanted me to do this but to be regarded as his botanical servant. He disliked changes in nomenclature that I proposed, including a new name. P. clusii, for P. cretica Tausch non Sabine, and the description of the Rhodes peony (P. rhodia) as a species distinct from P. arietina, with which he considered it conspecific, and these disagreements, together with our various other commitments, ended our co-operation. Luckily Stern had the supreme good fortune to obtain the help of J. Robert Sealy of the Kew Herbarium who prepared a key to the genus, drew up the descriptions, searched the literature and listed the specimens in the Kew Herbarium. In 1946 the Royal Horticultural Society published Stern's fine volume A Study of the Genus Paeonia, with illustrations by Lilian Snelling and Stella Ross-Craig (Mrs J.R. Sealy). For this work I had provided much of the information in the chapter IV ‘History of Paeonia literature’ and Appendix II, ‘Bibliography’. Maps on pages 18, 20, 22, 30 and 32 were prepared by George Sidney Holland and me. From two ladies came £ 1000 towards the cost of publication. But for Stern's enthusiastic interest in the genus Paeonia none of us would probably have given it the same attention. A Study of the Genus Paeonia stands as a splendid and very useful monument to Stern's interest in peonies over so many years; it is fitting that a peony species, P. sterniana Fletcher (1959), also commemorates him.

An appreciative but critical review of Stern's Study by G. Ledyard Stebbins appeared in MadroÔo 9: 193-199 (1948). Stebbins himself had earlier and independently given close study to the cytogenetics and the morphological characters of the species and hybrids cultivated in the U.S.A. and found himself unable to accept Stern's assumptions as to the significance of polyploidy and the relationship between taxa. In 1939 he published "Notes on some systematic relationships in the genus Paeonia" in University of California Publications in Botany 19: 2 15-266, which is of special interest for directing attention to the character of the sepals (see p. 49). Stebbins had no direct acquaintance with any Greek peonies which, however, well exemplify the advanced type of calyx with an abrupt transition from leafy outer sepals with a broad base and vestigial blade to large rounded inner sepals having no evident midrib. No Greek peony has a calyx with a gradual transition from elongated outer sepals to small inner sepals with evident midrib and a small apical point such as characterizes more primitive many-flowered Asiatic species, for example P. anomala and P. lactiflora.

The account of Paeonia by J. Cullen and V.H. Heywood in Flora Europaea 1: 243-244 (1964) recorded three species from mainland Greece and Crete, namely P. peregrina, P. clusii and P. mascula. Davis and Cullen in Flora of Turkey 1: 204-206 (1965) recorded from the East Aegean Greek islands P. mascula subsp. arietina from Samos and P. rhodia from Rhodes. None made use of the material in Greek herbaria, to which numerous specimens have been added since 1964. Indeed the collections now available, made in particular by Elli Stamatiadou and Dimitrios Tzanoudakis, have established the general patterns of distribution so well that further collecting is unlikely to modify these greatly (Figs 22, 23).

The results of combined cytological and taxonomic study based on extensive field work and collecting by Tzanoudakis are given in his thesis Cytotaxonomic Study of the Genus Pueonia L. in Greece (Patras, 27 April 1977). It is regrettable that this detailed study was not published fully in English, French or German, instead of only in Greek with a very concise English summary, as it makes a valuable contribution to understanding of the Greek peonies. Tzanoudakis studied cytologically 34 populations. The individuals of each population were either diploid (2n:2x:20) or tetraploid (2n:4x:40). Diploid as well as tetraploid populations were found in P. mascula and P. clusii, whereas all populations of P. peregrina and P. parnassica were tetraploid. However diploid plants of P. peregrina have been found in Jugoslavia which do not differ morphologically from the tetraploids. In this work Tzanoudakis described the peony with dark red flowers long known from Mount Parnassos as a new species, P. parnassica, and the white-flowered peonies of the P. mascula complex as subsp. hellenica and subsp. icarica. In the present work we have accepted these taxa mostly at the ranks assigned by Tzanoudakis but have maintained P. rhodia as specifically distinct from P. clusii and have united P. mascula subsp. hellenica and subsp. icarica.

CHAPTER II

GENERAL MORPHOLOGY content

Whether Paeonia is included in the Ranunculuceae, which has long been customary, and there treated as the only genus of the tribe Paeonieae, in accordance with Bentham and Hooker's Genera Plantarum 1:10 (1862), or associated with Glaucidiuin and Hydrastis, in accordance with Prantl in Engler and Prantl’s Die Natürlichen Pflanzenfamilien III., 2: (1888), or placed by itself in a family Paeoniaceae as proposed by Rudolphi and by Bartling in 1830 and accepted in the Flora Europaea 1:243 (1964), Hutchinson's Families of Flowering Plants, 3rd ed. 493 (1973) and most other modern works or associated in Paeoniaceae with Glaucidium as advocated by Melville (1983), its marked distinctness from other genera remains indisputable. The retention of primitive features, particularly in the Eastern Asiatic section Moutan, together with a wide distribution across the northern hemisphere, also indicates its great antiquity. Distinctive characters of the genus include a ring of vascular bundles in the stem, scalariform vessels, alternate pinnately or ternately compound leaves, unequal persistent sepals, large bowl-shaped corolla, very numerous stamens with marginally dehiscent anthers, tricolporate finely reticulate pollen grains, 1 to 6 free carpels with numerous ovules in two rows and large follicles with broadly ellipsoid or almost globose seeds having much endosperm and a small embryo. The development of the embryo is unusual (cf. Yakovlev & Yoffe, 1957; Carniel, 1967).

All the Greek peonies are herbaceous. They die down completely for part of the year and perennate by large buds on a subterranean root-stock from which descend fleshy roots providing nutriment for renewal of growth. The roots (Figs 15, 16) may be tapering, as in P. mascula, P. clusii and P. rhodia, or fusiform to subglobose as in P. officinalis, P. parnassica and P. peregrina, in which a slender string-like attachment growing from the rootstock becomes gradually or abruptly swollen, then narrows into a slender portion bearing the root hairs with the tubers hanging as it were on strings or very thick threads as in Filipendula vulgaris. This important distinction was well-known to the early her balists and portrayed in their works (Figs. 5-8) but ignored by most botanists because, of course, it can only be observed when the plants are uprooted, which is undesirable both in the garden and in the wild.

At the base of the stem are several sheaths sometimes regarded as stipules but probably best compared to modified petioles. The flowering stem always bears several long-stalked alternate leaves without stipules. Excluding the reduced leaf which sometimes occurs immediately below the calyx and counting only leaves distinctly separated from it, these vary in number from 3 to 12, but for each taxon a certain number of leaves predominates. Thus P. tenuifolia has usually 9 to 11 leaves, P. mascula subsp. russi usually 4 to 6. The variation cannot be ascertained from the limited number of specimens, often incomplete, in herbaria and hence requires study in the field to ascertain its value. Of 94 individual flowering stems of P. mascula subsp. hellenica examined on Andros, 50% had 5 leaves, 25% 4 leaves, 20% 6 leaves, with a range between 3 and 7. Of 56 flowering stems of P. peregrina examined on Lefkas, 34% had 8 leaves, 25% 9 leaves, 25% 7 leaves, with a range from 6 to 12. Of 48 flowering stems of P. mascula subsp. russi on Lefkas, 31% had 5 leaves, 29% 6 leaves, 25% 4 leaves, with a range from 3 to 7.

The leaves are ternately or pinnately compound. The basic type of leaf, as found in P. coriacea, is biternate; it consists of a long common petiole from which arise three primary petiolules, each bearing a primary division consisting of three primary segments or leaflets, thus giving 9 ultimate segments, one central, two lateral; sometimes a primary segment may be deeply two-or three-lobed. From this a more complicated leaf may be developed, by each of the central primary segments being itself divided into three secondary segments, thus producing a leaf with 16 ultimate segments. A further division of the three secondary segments, the central segment deeply and ternately divided, the lateral segment pinnately divided, produces an elaborately dissected leaf as in P. clusii, the whole possibly having as many as 80 ultimate lobes. An unpublished description of the leaf of P. clusii and a diagram (Fig. 17) both made about 45 years ago will illustrate the difficulty of portraying such a leaf in words: "Leaves biternate, glabrous (or very rarely pubescent beneath with sparse hairs), up to 38 cm long, 30 cm broad; blade finely cut with about 33 ultimate lobes; primary divisions 3, the central division long-petiolulate and divided into 3 primary segments cuneate at base with the central one deeply and ternately divided and gradually attenuate at base into a long petiolule, the lateral primary segments deeply and pinnately divided and sessile, the lateral divisions short-petiolulate (with the petiolule to 7 cm long) divided into 3 primary segments cuneate at base, the central one deeply and pinnately divided and gradually attenuate at base into a long petiolule, the outer lateral assymetrically 3-lobed, the inner one entire or slightly toothed towards the apex; ultimate lobes narrowly obovate to narrowly lanceolate, acuminate, from 5 cm long 2 cm broad to 5 mm long, 3 mm broad; common petiole (to 17 cm long) and petiolules cerise-tinged." From this it will be evident that the description of a leaf of a peony is not only complicated: its details, however essential, give no mental picture of the leaf as a whole. For Paeonia, as for other genera with much divided leaves, good illustrations are much more informative.

The leaves, however, provide the most important characters for the distinction of the European taxa even though the differences are not always clear-cut and in the herbarium may be hard to discern; indeed it is sometimes difficult to find two leaves exactly alike. The ultimate segments or lobes of the leaf, i.e. those at the end of its divisions and most remote from the petiole, vary from the broad almost orbicular or obovate leaflets of P. mascula subsp. triternata to the very narrow linear segments of P. tenuifolia, neither of which occurs in Greece. In the Greek species the ultimate lobes are mostly ovate or obovate to narrowly lanceolate and are glabrous to shortly pubescent on the underside. The petioles and petiolules are green in most species but red or reddish in P. mascula subsp. russi, P. clusii and P. rhodia. The uppermost leaf even though some distance below the calyx may be reduced to a single entire leaflet in P. parnassica as also in the Caucasian P. wittmanniana. The height of the (lowering stem is partly conditional on place of growth but is usually between 25 and 35 cm in P. mascula and P. clusii and between 60 and 80 cm in P. peregrina. The shrubby Asiatic species, such as P. suffruticosa and P. lutea, which are cultivated in European gardens, rise to about 1.5 metres.

The sepals (Fig. 18) show a marked change in form from the outside inwards. Immediately below the calyx there is often a small leaf with pinnate venation and a distinct midvein which may be divided or entire but is narrowed to the base; then usually come several progressively shorter sepals broad at the base and with an acute tip, these forming an involucre; then abruptly distinct from them in our Greek species, come the inner strongly concave and almost orbicular or very broadly ovate rounded sepals covering the petals and ofwhich the several veins arise directly from the base without a prominent midvein. This is characteristic of solitary-flowered peonies. Although there are some differences in the calyx between different species, these are difficult to define. In the Asiatic species with several-flowered stems, such as P. lactiflora and P. anomala, there is a gradual transition from the outer to the inner sepals and the midrib of the innermost sepals reaches the apex as emphasized by Stebbins (1939).

The open corolla may be markedly bowl-shaped, at first subglobose, with strongly concave petals, as in P. peregrina, to saucer-shaped with more widely spreading petals, as in P. mascula subsp. russi, and at anthesis about 7-13 cm broad. The colour in wild populations shows very little variation indeed and thus is of high taxonomic value. P. mascula subsp. hellenica, P. clusii and P. rhodia always have white flowers, occasionally with a faint pink tinge towards the base. P. mascula subsp. russi and others have rose-pink or purplish flowers, but in P. peregrina, the most spectacular of herbaceous peonies, they are a deep rich glossy red. P. parnassica has very dark red flowers.

The stamens are very numerous, about 140, with slender filaments, which may be white or rose, and yellow anthers dehiscing marginally, with very abundant yellow pollen. In the garden the pollen attracts many bees which make fleeting visits and rarely touch the stigma. In the wild, however, the main visitors appear to be large beetles which crawl among the stamens and over the stigma eating pollen, of which there is more than enough for all. Peony flowers are particularly suited for beetle-pollination: they provide a broad landing area, have no complexity, often emit a somewhat fusty smell and proffer their pollen lavishly. Since both beetles and peonies are of great antiquity, they have probably had a close association all through their history (cf. Leppik, 1964; Gottsberger 1977).

The pollen grains are spheroid prolate, 3-colporate, with the furrows (colpi) narrow and extending almost to the poles, the exine finely reticulate, 25-40 micro-m in length (Figs 19, 20).

Between the stamens and the carpels is a fleshy outgrowth, the disc, which is relatively inconspicuous in the Eurasian herbaceous species but much more developed in the Eastern Asiatic section Moutan and the North American section Onaepia.

The carpels are free to the base and range in number from one (which is normal in the North Indian P. emodi but only occasional in European species) to 6. The predominant number tends to differ from taxon to taxon. Thus in a population of P. mascula subsp. hellenica on Andros, with a 1-5 carpel range, 88 flowers being examined, 60% had 3 carpels, 17% 2 carpels, 14% 4 carpels, thus with an overwhelming predominance of 3 carpels. P. mascula subsp. russi, of which 56 flowers were examined on Lefkas, likewise with a 1-5 carpel range, had 32% with 3 carpels, 29% 2 carpels, 25% 4 carpels, thus with almost equality of 2, 3 and 4 carpels. On cultivated P. peregrina, 6 flowers had 1 carpel, 53 had 2 carpels, 9 had 3 carpels; of 9 flowers examined on Lefkas, 8 had 2 carpels but only one had 3; thus in P. peregrina flowers with 2 carpels appear to represent about 78% and 3 carpels 13%. In some Mediterranean and Asiatic peonies the carpels are glabrous although the plants as a whole may closely resemble others with densely hairy carpels. Stapf, quoted above from Curtis's Botanical Magazine 155: t. 9249 (1931), held that this difference was of high taxonomic value, entitling each member of the pair to specific rank, whereas most botanists have given them lower status as varieties or subspecies. Its value clearly depends on the extent to which it is correlated with other differences, including geographical distribution. In 1937 I asked Miss J. Coote to note whether there were any differences in appearance, ecology and distribution in Kashmir between plants of P. emodi with hairy carpels (var. emodi) and glabrous carpels (var. glabrata); she found none and observed that both occurred together. This problem does not arise in the classification of the Greek species. All have densely tomentose carpels, with glabrous sessile flattened recurving stigmas, which develop into tomentose follicles (Fig. 21). These vary somewhat in size. P. arietina derived its specific epithet from mature follicles being curved backwards somewhat like a ram's horns.

The fertile seeds are black or very dark brown, very broadly ellipsoid, with a glossy very minutely and densely pitted surface. They are 8-9 mm long, 6-7 mm broad in P. mascula subsp. mascula, hellenica, and russi when fresh and jet black, but slightly larger (8-10 mm long, 7-8 mm broad) and brownish black or jet black in P. peregrina.

The characteristics of the fruits were aptly described by the English herbalist John Gerard in 1597, who stated that when the ‘faire large redde floures very like roses' have fallen away there come in place three or foure great cods or husks, which do open when they are ripe; the inner part of which cods is of a faire red colour, wherein is contained blacke shining and polished seeds, as big as a Pease, and between every blacke seed is couched a red or crimson seed, which is barren and empty.”

CHAPTER III

GENERAL DISTRIBUTION content

Paeonia is a genus of the northern hemisphere divisible into three sections: Paeonia (Paeon DC.), with P. officinalis as lectotype, Moutan DC., typified by P. suffruticosa, and Onaepia Lindley, typified by P. brownii. Onaepia with two species is confined to western North America, Moutan to eastern Asia. The section Paeonia, to which belong all the Balkan peonies, extends from Spain eastwards to China and Japan, spills over into north-west Africa and just reaches the Arctic Circle in the Kola Peninsula. The genus is basically one of light woodland on limestone slopes although in the Mediterranean region some populations survive in degraded macchie and among rocks. Individual plants are deep-rooted and long-lived but are nevertheless vulnerable and everything possible should be done to prevent their extinction.

Peony seeds are smooth and relatively large. They lack any obvious means of dispersal. The present distribution of the Greek peonies (Figs 22-24) suggests that they attained their present range by very gradual spread over a very long period of time. Long distance dispersal has probably had no part in this although regression of range through local extinction presumably has. They now occur in widely separated areas. A peony first described from Sicily as P. russi (now P. mascula subsp. russi) is now known from the Ionian Islands in the Adriatic Sea and adjacent mainland (prov. Akarnania); this has purplish red (mauve) flowers. A peony with white flowers (which become yellowish on drying) has long been known from the northern mountains of Sicily and was named P. flavescens by C.B. Presl in 1822. It does not, however, appear to be separable from P. mascula subsp. hellenica which occurs in the Taygetos mountains of the Peloponnese and is there far distant from a colony in Attica, which in turn is remote from those on Euboea (Evvia), Andros and Ikaria. The affinities of most Greek peonies are with plants of Asia Minor. The only endemic species are P. clusii on the islands of Crete and Karpathos and P. rhodia on Rhodes.

The distribution of these peonies reflects changes long ago in the distribution of land and water in the Aegean region. Flans Runemark (1971), using data from Creutzburg (1963, 1966) has outlined the relevant palaeography of the Aegean as follows (very slightly paraphrased): "The south island chain is part of the Alpine fault system. The islands of Rhodes, Karpathos and Crete have been isolated from each other and from the mainland for 3-5 million years. In the Pliocene a land-bridge existed in the central part of the Aegean, connecting Southern Greece and Western Turkey. It was delimited to the North by a large lake, covering most parts of the present North Aegean Sea, and in the South by the Sea of Crete, which separated it from the South Aegean island chain. Connections with the latter never seem to have existed. "The first fault zone separating the land-bridge from Turkey was established in the beginning of the Pliocene... between Astypalaia and Kalimnos, and Ikaria and Samos. Soon Ikaria seems also to have been isolated from the rest of the Central Aegean. Comparatively later the Central Aegean was permanently separated from the Greek mainland by a channel between Attica and Kea and between Euboea and Andros. The central land mass was afterwards split, resulting in the present archipelago. Most large islands in the Aegean have apparently been isolated for hundreds of thousands of years... During the Pleistocene, the sea level fluctuated considerably with regressions up to 120 m — according to Pfannenstiel (1951, 1954) up to 200 m — during glacial periods and transgressions up to 35 m during interglacials."

These physical changes left the white-flowered peonies in isolated but logical positions. Paeonia mascula subsp. hellenica (2n:20) confined to Attica (where it is now very rare) and the islands of Euboea (Evvia) and Andros, with an outlying station in Laconia in the southern Peloponnese, and a colony (var. icarica) on the schists of Ikaria. Separated by the persistent Sea of Crete, the white-flowered P. clusii (Crete and Karpathos; 2n:10, 20) and P. rhodia (Rhodes; 2n:10) remained confined to the South Aegean island chain. Reference to the distribution maps (Figs 22-23) will demonstrate their present occurrence.

The present distribution of the red-flowered peonies in mainland Greece has a different but probably simpler explanation than that of the very localised white-flowered taxa on the islands. The red Greek peonies are part of a wider distribution pattern in Europe and the Near East. P. peregrina (2n:20) in northern and western Greece has a somewhat northerly Mediterranean distribution that extends from Italy to Turkey. P. parnassica (2n:20), now confined to the vicinity of Mt. Parnassos, is presumably an isolated derivative of P. peregrina. P. mascula is a collective species with well distinguished subspecies. Subsp. mascula (2n:20), though localised in Greece, is part of a European distribution that extends from central France across central and southern Europe to Turkey, Cyprus, Caucasia and northern Iran. Plants with some characters of subsp. triternata (P. daurica; 2n:10) occur very locally in the eastern Aegean islands of Greece. Typical subsp. triternata possibly occurs only in the Crimea but plants related to it have a disjunct distribution in Jugoslavia (Bosnia), northern and southern Anatolia, Transcaucasia and northern Iran. Subsp. russi (2n:10), a diploid, was first recognised in Greece by Tzanoudakis (1977), from the Ionian Islands and western Greece, an eastern extension of a disjunct island range on Corsica, Sardinia and Sicily; indeed, it might be considered a relict Central Mediterranean subspecies (Fig. 24).

Tzanoudakis's Cytotaxonomic Study of the Genus Paeonia in Greece (1977) provides a critical assessment of the origin and relationships of the tetraploid peonies. Here he states that "According to the cytological data, the tetraploids of P. clusii (eastern Kriti) should be considered as autotetraploids derived from diploids of the same species; such diploids occur today in western Kriti and Karpathos. On the contrary, the tetraploids of P. mascula, P. parnassica and P. peregrina should be considered as allopolyploids derived as follows: a) Those of P. mascula (subsp. mascula, subsp. icarica and subsp. hellenica) from a cross of P. mascula subsp. russi with P. clusii or from a cross between their ancestors or their autotetraploid derivatives, b) Those of P. parnassica from a cross between autotetraploid derivatives of P. mascula subsp. russi and P. peregrina. The origin of P. peregrina remains problematic; although cytologically it resembles an allopolyploid, diploid plants have been discovered in Jugoslavia with no morphological differences from the tetraploids (Sopova 1971)."

The lack of discernible differences between peony populations of the same subspecies which have been isolated over very long periods of time exemplifies "the evolutionary standstill which affects the bulk of the Aegean relict species" noted by Greuter (1979).

CHAPTER IV

THE GREEK SPECIES content

Paeonia L., Sp. Pl. 1:530 (1753), Gen. Pl. 5th ed. 235 (1754).

Lectotype: P. officinalis L. sensu stricto (var. [alpha] feminea).

Perennial herbs. Leaves alternate, at least the lower ones simply biternate or further divided. Flowers solitary, large. Sepals 3-8, imbricate, herbaceous, persistent. Petals 6-9, broad, red, pink or white. Stamens numerous, maturing centrifugally. Carpels 2-6 (-8), multiovulate, surrounded at base by a depressed fleshy disc. Styles with a circinnate stigmatic area. Fruit a group of thick follicles with numerous seeds; fertile seeds blackish, borne on a ± fleshy funicular aril; sterile seeds smaller, red and juicy.

The above description applies particularly to the Greek species, all of which belong to the section Paeonia (Paeon DC.), not to the genus as a whole for which see Cronquists's description (1981).

The standard abbreviations for names of herbaria are used in the lists of specimens testifying to geographical distribution: BM, British Museum (Natural History), London; ATH, Goulandris Natural History Museum, Kifissia; ATHU, Department of Botany, University of Athens; E, Royal Botanic Gardens, Edinburgh; K, Royal Botanic Gardens, Kew; UPA, Department of Botany, University of Patras. In accordance with botanical custom over more than 150 years, the exclamation mark(!) following such an abbreviation indicates that a specimen of a particular gathering has been examined in the herbarium cited. Thus “(ATH! BM! K!)” indicates that specimens have been examined in the Kifissia, British Museum and Kew herbaria, but a plain citation, e.g. “(UPA)” indicates that the specimen at Patras studied by Tzanoudakis has not been seen during the preparation of this work though accepted on his authority. Localities are listed in the sequence of Greek administrative provinces and districts enumerated by Greuter (1977).

KEY TO GREEK SPECIES OF PAEONIA content

1. Lower leaves with obovate-cuneate leaflets, furcate or shortly lobed; flowers dark red ....................... 1. peregrina

1. Plants not as above ...................... ... 2

2. Stem hirsute; lower leaves biternate with elliptic to lanceolate leaflets profusely pilose beneath; sepals pilose; petals dark red 2. parnassica

2. Stem glabrous or nearly so; lower leaves glabrous or pubescent beneath, with 9-80 leaflets; sepals glabrous; petals rod, purplish, pink ......... 3

3. Lower leaves basically biternate, with 9-21 leaflets, glabrous or pubescent beneath; petals red, pink or white ............. 3. mascula

3. Lower leaves with 9-80 leaflets or lobes, glabrous; petals white. .................. 4

4. Lower leaves with 40-80 leaflets and lobes, the leaflets usually 3-5-fid to 1/2 or 2/3 into oblong-lanceolate lobes ............... 4. clusii

4. Lower leaves with 9-29 leaflets and segments, mostly ovate to oblong-elliptic or lanceolate ..................... 5.rhodia

The above key includes all the Balkan and Aegean species except P. officinialis L. (1753), illustrated here on pages 24, 27, 28 and 30, and P. tenuifolia L. (1759) easily recognized by its intricately divided leaves with very numerous linear segments not more than 5 mm wide. P. officinalis occurs in northern Jugoslavia and Romania, P. tenuifolia in Jugoslavia, Romania and Bulgaria, but neither in Greece.

1. P. peregrina content Plate 3; Figs 12, 13,21,25

Paeonia byzantina Clusius, Har. PI. Hist. 279, fig. (1601).

P. peregrina Miller, Gard. Dict. ed. 8, art. Paeonia No. 3 (1768); Stapf in Curtis's Bot. Mag. 144: t. 8742 (1918): F.C. Stern. Study of Paeonia 97 (1946); Nyárády in Savulescu, Fl. Republ. Pop. Romane 2:404 (1953); Cullen & Heywood in Fl. Europ. 1:243 (1964); Davis & Cullen in Fl. Turkey, 1:206 (1965); Goulandris, Goulimis & Stearn, Wild Flowers of Greece, 22 t. (1968); Jordanov, Fl. Reipubl. Pop. Bulgar. 4:219, t. 42 (1970); Tzanoudakis, Cytotaxon. Study of Paeonia in Greece, 39 (1977).

P. decora G. Anderson in Trans. Linnean Soc. London 12:73 (1818); Hayek, Prodr. Fl. Penins. Balcan. 1:298 (1924).

Illustrations: Sweet, Brit. Flower Garden, 1: t. 70 (1824) as P. lobata; Curtis's Bot. Mag. 144: t. 8742 (1918); Goulandris, Goulimis & Stearn, op. cit. 22 (1968); Jordanov, op. cit. 4: t. 42 (1970).

Protologue: “3. PAEONIA (Peregrina) foliis difformiter lobatis, lobis incisis, petalis florum rotundioribus. Peony with deformed lobated leaves which are cut, and rounded petals to the flower. Paeonia peregrina, flore saturate rubente. C.B.P. 324. Foreign Peony with a deep red flower. The third sort grows naturally in the Levant; the roots of this are composed of roundish knobs like those of the second sort [P. officinalis senso stricto, P. feminea], as are also the leaves, but are of a thicker substance: the stalks do not rise so high, and the flowers have a greater number of petals. This flowers a little after the other." (Miller, loc. cit. 1768).

Description: Stem glabrous, 30-50 cm. Roots slender then swollen into cylindric or fusiform tubers. Lower leaves biternate, firm, with some leaflets divided to base, in all with 15-17 principal divisions, some of these deeply cut into 2-3 segments; segments and divisions coarsely toothed at apex; principal divisions 5-12 cm long, glossy green above, glaucous, glabrous or sparsely pilose beneath. Flowers 7-12 cm across, strongly concave. Petals 6-7 (-10), oblong-ovate to suborbicular, deep red, often split when pressed. Filaments pink or red. Carpels 1-4, mostly 2, long tomentose. Style c. 9 mm; stigmatic area circinnate from near base, c. 2 mm broad. Fl. April - May. 2n: 20.

Type: Described from cultivated material originally introduced from “the Levant”, i.e. from Istanbul in the late 16th century (Fig. 11).

Distribution: Ionian Islands, north-eastern and northern Greece, with isolated occurrence in eastern central Greece, and island of Thasos.

THRAKI (Thrace). Prov. 11, Evros: district Alexandroupolis, ravine W. of Alexandroupolis opposite of Avra, iv. 1951, Goulimis 16030 (ATH!) distr. Alexandroupolis, 11-12 km from Loutros to Pessani, 260-300 m, Stamatiadou, 15209 (ATH! BM!); district Soufli, in Pessanis forest, v. 1961, Goulimis 16031 (ATH!).

ANATOLIKI MAKEDHONIA (Eastern Macedonia). Prov. 21, Kavala: district Thasos, 400 m, Dimonie (G). Prov. 24, Halkidhiki: Ajion Oros (Mount Athos) near Chalandari, Guiol (BM!).

DHITIKI MAKEDHONIA (Western Macedonia). Prov. 37, Imathia: prope pagum Gianakochori, Tzanoudakis 2337 (UPA).

IPIROS (Epirus). Prov. 51, Ioannina, prope pagum Grammeno, c. 500 m., Tzanoudakis 2222 (UFA).

IONII NISI (Ionian Islands). Prov. 62, Lefkas: Mount Stavrotas, north of Agios Ilias, 900-1150 m. Stamatiadou 12240 (ATH! BM!).

STEREA ELLAS (Central Greece). Prov. 73, Fthiotis (Phthiotis): district Dhomokos, Palamas, Dhramata, v. 1957, Goulimis 16034 (ATH!).

Greek habitats: Quercus forest, in dry and in damp places among limestone rocks, dry rocky ground with shrubs. Altitude 200-1150 m.

External Distribution: Italy, Jugoslavia. Albania, Bulgaria, Romania.

Paeonia peregrina is the most gorgeous of Greek peonies when the sun lights up its large bowl-shaped ruby red flowers and it is also the most distinctive in foliage. It was introduced into cultivation in Austria in the late 16th century by way of Constantinople, as recorded by Clusius in 1583, and passed from garden to garden across Europe; by 1629 Parkin-son grew this "single red peony of Constantinople" in England. Miller's name for it, P. peregrina, published in 1768 was misapplied by most later authors to other species until in 1899 Carl Fritsch brought that name back into use for this Balkan species which meanwhile had been named P. decora and P. romanica. Records of P. peregrina from Italy mostly do not refer to this species, but a specimen collected by C.C.Lacaita in Calabria Citra, Monti di San Donate di Ninea, at 1200 m., in 1912 (BM!) testifies to its occurrence, apparently very rare, in southern Italy. It received the epithet peregrina, meaning ‘exotic, foreign, coming from abroad,’ early in the 17th century because it was then a newcomer to Central European gardens unlike P. officinalis and P. mascula.

2. P. parnassica content Plate 4; Fig. 26

P. peregrina var. latifolia Boiss., Fl. Orient. 1:97 (1867).

P. parnassica Tzanoudakis, Cytotaxon. Study Paeonia in (Greece, 43 (1977); Broussalis in Physis (Athens) 14:13, fig. (1978).

Illustration: Broussalis, loc. cit. (1978).

Protologue: "4. P. parnassica Tzanoudakis, sp. nova, Typus: Prov. Phokis, mons Parnassos, supra pagum Agoriani, ca 1100 m, in Abietis Tzan. 1400 (Hololypus UPA). Syn P. peregrina ([gamma]) latifolia Boiss., Fl. Or. 1:97 (1867). P. arietina auct." (non Anderson in Trans. Linn. Soc. London 12:275, 1818).

"Planta perennis. Radicis tubera numerosa, fusiformia nec non sub-globosa. Caulis dense pubescens, florifer uniflorus, viridis, (30) 35-40 (50) cm altus. Folia biternata subtus dense pubescentia, segmenta secundaria elliptico-ovata vel lanceolata, integra vel lobata, 4-8 cm longa et 2-4 (6) cm lata. Flos c. 8-10 cm diametro. Petala atropurpurea, obovata vel late obovata. Sepala 3-5, plerumque aequalia, pubescentia, obtusa usque ad mucronata, 1-2 cm lata. Carpella 1-2 (3), tomentosa, stigmatibus sessilibus. Planta tetraploidea, chromosomatibus 20.

Prov. Phokis: Supra pagum Agoriani, Pinatzi 17782 (Herb. Pinatzi), Tzan. 1400 (UPA). In monte Parnasso prope Carcaria, Orph. 445 (ATHU). In monte Parnasso, supra Arachova ad Ag. Nikolaos, Heldreich 2648 (ATHU). Prov. Boeotia: Mons Elikon, in regions Abietina, Pinutzi 203 (Herb. Pinatzi)." (Tzanoudakis, loc. cit., 1977).

Description: Stem 35-65 cm, hirsute. Lower leaves biternate; lateral divisions with the terminal leaflet entire or deeply 2-3 lobed; leaflets and their lobes 9-13 in all, obovate to narrowly elliptic or lanceolate, acute or shortly acuminate, cuneate to attenuate at base; leaflets green above (purplish when young), greyish green and densely pilose beneath; terminal petiolules of terminal leaflets 5-15 cm. Flowers 8-12 cm across. Petals 9-12, obovate to obovate-orbicular, dark red. Filaments purplish. Carpels 2-3, tomentose. Style circinnate and stigmatic almost from base, c. 1,5 mm broad. Fl. May 2n : 20.

Type: Greece. “Prov. Phokis, mons Parnassos, supra pagum Agoriani. c. 1100 m.. in Abietis Tzan. 1400” (UPA, holotypus; BM!)

Distribution: Endemic to mountains of southern central Greece.

STEREA ELLAS (Central Greece). Prov. 74, Fokis (Phocis): distr. Parnassis: Parnassus prop. Carcaria, 5000 ped., 1854, Orphanides (BM! K! type-collection of P. peregrina var. latifolia Boiss.); Karkaria. 1863, J.S. Mill (K!); 1969, Merkniia (ATH!); 16-17 km S.E. or village Eptalofos, in valley 1-1.5 m N.N.W. of Agios Nikolaos, 1200-1300 m, 23.V.1982, Stamatiadou 22045 (ATH! BM! E!).

Prov. 75, Viotia (Boeotia): Mount Helikon, 4000 ft., v-vi. 1934, Atchley 200 (K!).

Habitat: Margins and openings in Abies cephalonica forest, damp grassy meadows and sheltered places between limestone rocks on south-western slopes. Altitude 800-1300 m.

Although known from herbarium specimens collected on Mt. Parnassos since 1854, P. parnassica was not recognized as distinct from P. arietina, under which the Kew specimens were listed in Stern's Study 81 (1946), until studied in a living state by Tzanoudakis and described in his Cytotaxonomic Study. It stands closest to P. peregrina but differs in leaf-form and in presence of hairs on the stem, on the underside of the leaf and on the sepals. The flowers are a dark blackish red unlike those of any other Greek peony, in colour more like those of the Chinese P. delavayi Franchet.

3. P. mascula content

P. officinalis L. var. mascula L., Sp. PI. 1 (1753). P. mascula (L.) Miller, Gard. Dict. ed. 8 art. Paeonia No. 1 (1768); Cullen & Heywood in Fl. Europ. 1:244 (1964).

Protologue: “1. Paeonia (Mascula) foliis lobatis ex ovatolanceolatis. Haller, Helv. 311. Peony with lobate leaves which are oval and spear-shaped. Paeonia folio nigricante splendido, quae mas. C.B.P. 323. Peony with dark shining leaves, otherwise male Peony. The first sort here enumerated, is the common male Peony, which grows naturally on the Helvetian mountains." (Miller, loc. cit. 1768).

Description: Very variable. Roots tapering, somewhat carrot-like. Stems 25-60 cm, glabrous. Lower leaves basically biternate; leaflets entire, or some of them deeply bifurcate or trifurcate; total number of leaflets 9-21, ovate, elliptic or broadly obovate, glabrous or pilose beneath, the terminal leaflets petiolulate. Flowers 7.5-13 cm across. Petals 5-9, red, pink or white. Filaments red, purple, pink or white. Carpels 2-5. Style 6-12 mm (including stigma). 2n : 10 or 20.

KEY TO SUBSPECIES

• 1. Petals white ..... 3 c. subsp. hellenica

• 1. Petals red, purplish or pink ...... 2

  • 2. Leaves purplish and not fully expanded at flowering time, long-pilose on lower surface. Stem 20-45 cm high. Follicles 1.5-2.5 cm long ..... 3b subsp. russi

  • 2. Leaves green and fully expanded at flowering time, glabrous or very sparsely hairy beneath. Stem 35-60 cm high. Follicles 2.5-4.5 cm long ....... 3

    • 3. Leaflets 9-12, flat, at least twice as long as broad, distinctly acute ............. 3a subsp. mascula

    • Leaflets basically 9 (a few divided at base in 1-2 small leaflets), undulate at margin and somewhat concave above, much less than twice as long as broad, obtuse or subacute ..... 3d. subsp. triternata

3a. P. mascula subsp. mascula content Plates 5,6,7; Figs 6,9,27

P. mascula (L.) Miller, loc. cit. (1768): Moss, Cambridge Brit. Fl. 3:155 t. 166 (192U); F.C. Stern, Study of Paeonia, 17, figs. (1946) in part P. corallina Retzius, Obs. Bot. 3:34 (1783); Sowerby & Smith, Engl. Bot. 22: t. 1513 (1806); Hayek, Prodr. Fl. Penins. Balcan. 1:297 (1924).

Illustrations: Sowerby & Smith, op. cit. t. 1513 (1806); Moss, op. cit. t. 166 (1920); F.C. Stern, op. cit. (1946).

Description: Stem 36-60 cm, glabrous. Lower leaves basically biternate, with 1 or several leaflets often deeply bifurcate or trifurcate so that the leaves have 9-21 leaflets. Leaflets nearly flat, elliptic to obovate-elliptic, broadly and shortly acuminate, green above, glaucous and glabrous beneath (rarely with few sparse hairs on veins); terminal leaflets broadly elliptic, flat, cuneate at base, attenuate into a 5-20 mm petiolute. Flowers 9-12 cm across. Petals 6-8, obovate, purplish red (fuchsia). Filaments purple. Carpels 2-5, long-tomentose. Style 8-12 mm; stigmatic area c. 2 mm broad, circinnate from base. Fl. April - May. 2n: 20. Type: Described from garden material (see below).

Distribution: Eastern central Greece and eastern Aegean islands.

STEREA ELLAS (Central Greece). Prov. 75, Viotia (Boeotia): supra pagum Koukoura, Tzanoudakis 1387 (UPA); supra pagum Elikon, Tzanoudakis 2257 (UPA). Prov. 76, Evvia (Euboea): supra pagum Rouklia, Tzanoudakis 1220 (UPA).

NISI TOU EJEOU (Aegean Islands). Lesvos (Lesbos): W.S.VV. of the Sanatorium of Ajiassos, 600 m, 17.v.1969, Stamatiadou 6309 (ATH!); S.W. of Ajiassos, on N.E. slope of Mt. Olymbos, below Profitis Elias Chapel, 750 m, 18.V.1969 Stamatiadou 6334 (ATH!). Samos: Mt. Ambelos, v. 1954, Goulimis 16029 (ATH!) Mt. Ambelos, 1-1.5 km S.S.W. of village Ambelos, on N. slope of Zovrahia summit 800-1000 m, 4.v. 1968, Stamatiadou 27336 (ATH! BM!). Mt. Ambelos, 1 km S.S.W. of Vronta monastery, on N.E. slope of Lazaros summit, 700-850 m. 29.iv.l970, Stamatiadou 8307 (ATH!); Ajiassos, 14.IV.1973 Broussalis (ATH!); Mt. Kerkis near Kakoperato, 900 m, Davis 1639 (E!).

Greek habitats: Shady pinewoods, damp shady places with Pteridium in Pinus-Abies forest, N.E. limestone slopes and clearings. Altitude 700-1200 m.

External distribution: Southern Europe eastwards to the Caucasus, Asia Minor, northern Iraq and Iran.

On Samos typical subsp. mascula and plants approaching subsp. triternata occur on Mt. Kerkis, whereas on Mt. Ambelos apparently only subsp. mascula is present. Stern (Study Gen. Paeonia. 81) records P. arietina Anderson (i.e. P. mascula subsp. arietina (Anderson) Cullen & Heywood) from Mt. Ambelos (Rechinger 3889) in Samos, but all material we have seen from this mountain has glabrous petioles etc., and is assignable to subsp. mascula. Subsp. arietina. however, grows in Anatolia and extends westwards to Kaz Dag (Trojan Ida) north of the island of Lesvos.

When separating as species the peonies hitherto known and distinguished metaphorically as male and female, Philip Miller rejected Linnaeus's 1753 comprehensive name P. officinalis but adopted his varietal epithets feminea (as foeminea) and mascula for specific epithets. Miller, as was his custom when using Linnaean epithets, did not directly cite Linnaeus's names as synonyms but cited pre-Linnaean polynominals also occurring in Linnaeus's Species Plantarum. Such common synonyms directly link Miller's Gardeners Dictionary, 8tb edition, and Linnaeus's Species Plantarum. The protologue of P. officinalis [beta] mascula L. (1753) is as follows:

"mascula [beta] Paeonia foliis lobatis ex ovato-lanceolatis Hall. Helv. 311. Paeonia folio nigricante splendido, quae mas. Bauh. pin. 323.

Paeonia mas. Lob. ic. 684.

Reference to Miller's protologue (cited above) shows that both he and Linnaeus cited the same works, Albrecht von Haller's Enumeratio methodica Stirpium Helvetiae indigenarum (1742) and Caspar Bauhin's Pinax Theatri botanici (1623); both authors took their diagnostic characters from Haller, whose account is as follows:

"2. PAEONIA foliis lobatis, ex ovato lanceolatis.

Paeonia mas Gesn. Hort. p. 270. Coll. p. 97. b. Matthiol. p. 914. Lob. ic. p. 684-685. Dod. p. 194. Tab. p. 784. Raj. p. 693. Blakw. T. 245.

Paeonia altera Trag. p. 582.

Paeonia Caes. p. 588.

Paeonia mas major flore incarnato, Aichst. Vern. Ord. VI T. XI. f.l.

Paeonia mas una praecocior I.B.III. p. 492.

Paeonia flore nigricante splendido quae mas C.B.P. I.R.H.

Paeonia simplex latiore folio trifido H.OX.III p. 454.

[beta] Paeonia mas altera tardior I.B.lc. H.O.Xlc. RAJ. p. 694 I.R.H C. Gesner in M. Generoso prope Luganum nasci audivit. Genevae Lobelius. Huic radix crassa, divaricata, non glandulosa, caulis altior, ad duos cubitos, ramis et costis foliorum purpura tinctis, fbliis amplioribus, minus difformibus, raro divisis, lobis tripinnis et quinquepinius, floris purpureo colore dulutiori, seminibus rotundis, crassioribus”

Both from the synonymy and the descriptive note on the roots (unlike those of his first species, P. officinalis “radicibus glandulosis interruptis") it is evident that Haller referred here to the Paeonia mas of earlier authors but did not know it as an undoubted Swiss native plant. In fact, unlike P. officinalis, it nowhere grows in Switzerland although widespread in Italy. Haller obviously knew it from living cultivated plants. L’Obel's woodcut (Fig. 9) on p. 684 of his Plantarurn seu Stirpium Icones (1581) cited by both Haller and Linnaeus clearly portrays the plant to which both Linnaeus and Miller applied the epithet mascula. Meikle, Flora of Cyprus 1:69 (1977) has suggested that it might be taken as lectotype of the name P. mascula (L.) Miller. Originally used in a narrow sense for a cultivated plant probably introduced from Italy, the name is now used to cover a diversity of recognizable taxa treated for convenience as subspecies but most of which have often been regarded with good reason as independent species.

3b. P. mascula subsp. russi content

Plates 2, 8, 9; Figs 14, 15, 28, 29

P. russi Bivona, Stirp. Rar. in Sicilia Descr. 4:12 (1816); Gussone, Fl. Siculae Prodr. 2:29 (1828-32); F.C Stern, Study of Paeonia, 63, fig.(1946).

P. corallina var. pubescens Moris, Fl. Sardoa 1:64, t. 4 (1837).

P. corallina var. russi (Biv.) Cosson, Comp. Fl. Atlant. 2:53 (1887).

P. revelieri Jordan in Jordan & Fourreau, Icones Fl. Europ. 2:38, t. 322 (1903).

P. mascula subsp. russi (Biv.) Cullen and Heywood in Feddes Repert. 69:35 (1964), Fl. Europ. 1:224 (1964); Tzanoudakis, Cytotaxon. Study of Paeonia in Greece, 34 (1977).

Illustrations: Moris, op. cit. t. 4 (1837); Jordan & Fourreau, op. cit. t. 322 (1903); F.C. Stern, op. cit. 64, 124 (1946).

Protologue: “POENIA RUSSI Peonia foliis biternatis, foliolis ellipticis, integris capsulis recurvatis pilosis, radice fusiformi. Radix fusiformis, simplex vel ramosa, ramis 1-2, pariter fusiformibus, patentibus. Caulis erectus, pedalis et altior, teres, subangulatus glaber, simplex, foliosus, purpurascens uti petioli. Folia inferiora biternata, summa saepe ternata, potiolo tereti, superne sulcato, foliolis obscure viridibus, ellipticis, integris, supra glabris, subtus vix pubescentibus. Calycis foliola quinque, quorum 2-4 parva, oblonga, concava, reliqua majora, elliptica, integra, raro biloba. Petala 5-6, obovata, chermisina. Capsulae duo, recurvatae, pilosae, pilis flavis, latere interiori dehiscentes.

Panormi in montibus cum Peon. officinalis et corallina. Perennis Floret Majo.

Observ. Paeoniae humili quam corallinae proximior, at differt ab illa forma radicis, integritate foliolorum et directione capsularum.

Dixi in honorem solertissimi patriae Botanices cultoris D. Joachimi Russo Cassinensis " (Bivona, 1816).

Description: Stem glabrous, 25-45 cm, purplish when young. Leaves purplish and not fully expanded when in flower; lower leaves biternate; leaflets ovate to obovate or broadly elliptic, flat, in fruit up to 10-12 cm long, 6-8 cm broad, apex shortly acuminate, blade densely to weakly pubescent over lower surface. Petiolule of terminal leaflets 8-15 (-20) mm. Flowers 9-12 cm across. Petals (5-) 6-8 (-9), obovate-orbicular, pinkish mauve. Filaments white or sometimes rose. Carpels 1-4, shortly tomentose. Style c. 6 mm long; stigmatic area 2 mm broad, undulate-circinnate from slightly above base. Fl. March - April. 2n: 10.

Type: Described from Sicily.

Distribution: Ionian Islands and western central Greece.

IONII NISI (Ionian Islands). Prov. 62 Lefkas (Levkas): Strongilo, E. of village Hortata, 900 m, 16. iv. 1970, Stamatiadou 8113 (ATH! BM! E!) places called Ptelia and Strongilo, 850-900 m., 20. iv. 1977, Stamatiadou 19652 (ATH! BM!). Prov. 63, Kefallinia (Cephalonia): Mt. Enos, S.W. slopes of Roudi summit, 700-800 m, 29. iv. 1974, Stamatiadou 17797 (ATH! BM!) Prov. 64, Zakinthos: on road from town to Anafonitria (Plemonaria) iv. 1952, Goulimis 16039 (ATH!); village Orthonies near the monastery of Spileotissis, 450-500 m, 13. iii. 1971, Merkatis (ATH! BM!); south of Spileotissis monastery, 400 m, 20. iii. 1976, Stamatiadou 18809 (ATH! BM! E!)

STEREA ELLAS (Central Greece). Prov. 71, Etolia - Akarnania: district Vonitsa- Xiromeros, Mts. Akarnanika, S. of the village Komboti, on E.S.E. foothills of Boumistos summit, 700-750 m, 28. vi. 1970, Stamatiadou 10315 (ATH!); 700-1000 m, 11. v. 1974, Stamatiadou 17845 (ATH!).

Greek habitats: Rocky limestone slopes with Quercus coccifera, Crataegus, Phlomis fruticosa between macchie zone and Abies cephalonica forest, and by vineyards. Altitude 400-900 m.

External distribution: Corsica, Sardinia, Sicily. The two gatherings from Sterea Ellas, Stamatiadou 17846 and 10315 (ATH), differ from typical subsp. russi in having a slender style recurved and stigmatic only in the upper third.

Two peonies occur in the mountains of Sicily, both now placed as subspecies in the collective species P. mascula but well distinguished: one mauve-flowered with leaves slightly pubescent below (subsp. russi), the other white-flowered and glabrous (subsp. hellenica). The mauve-flowered one was known to Francesco Cupani (1657-1711) but was first named, as P. russi, by Bivona-Bernardi in 1816. This and Saxifraoa russi C. Presl (1822) commemorate the abbé Giocchino Russo from Monte Cassino, who was evidently keenly interested in the flora of Sicily during the first quarter of the 19th century. As his name would have been latinized as Russus, Bivona-Bernardi and Presl both used the genitive epithet Russi. This was first recorded from Sicily as Paeonia hyemalis, pumila, Rosae rubrae monoflore in Cupani's Hortus Cathiolicus 169 (1696) and later illustrated by an engraving (Fig. 14) in his extremely rare and almost always incomplete Panphyton Siculum (1713).

3c. P. mascula subsp. hellenica content

Plates 10, 11, 12; Figs 30, 31, 32

P. flavescens C. Presl in J.S. & C. Presl, Deliciae Prag. 5 (1822); C. Presl, Fl. Sicula 27 (1826).

P. corallina var. flavescens (C. Presl) Gussone, Fl. Siculae Prodr. 2:28 (1828-1832), Fl. Siculae Syn. 2:26 (1844); Lojacono Pojero, Fl. Sicula 1:53 (1888).

P. mascula var. flavescens (C. Presl) Gürke in Richter, Pl. Europ. 2:400 (1903).

P. mascula subsp. hellenica Tzanoudakis, Cytotaxon. Study Paeonia in Greece, 36 (1977).

P. mascula subsp. icarica Tzanoudakis, op. cit. 38 (1977).

a) var. hellenica

Protologue: "P. mascula subsp. hellenica Tzanoudakis subsp. nova.

Typus: Insula Euboea, Kerasovrysi, in ditione pagi Steni, Tzan. 1210 (Holotypus UPA).

A subspeciebus mascula et russi floribus albis, maioribus differt; a sub-specie icarica foliis subtus minus pilosis, segmentis secundariis segmenti centralis primoris indivisis differt.

Prov. Attica: Mons Parnis, Sotiropoulos 252 (UPA).

Ins Euboea, Kerasovrysi in ditione pagi Steni, Phitos 10335 (UPA), Tzan. 1210 (UPA), Pinatzi 11882 (Herb. Pinatzi). Mons. Delfi, supra Steni, Orphanides 472 (ATHU). In mont. Xiron oros, Rechinger 17105 (G), Pinatzi 202 (Herb. Pinatzi). Supra pagum Metochi Dirphios, Tzan. 1205 (UPA).

Mons Ochi, supra pagum Rouklia, Tzan. 1220 (UPA).

Ins. Andros; supra Arna, Tzan. 1954 (UPA).

Prov. Laconia: Mons Taygetos, Goulimis 7373 (Herb. Pinatzi). Supra pagum Spartia, Tzan. 1286 2347 (UPA).“ (Tzanoudakis, loc. cit. 1977).

Protologue: “2d. P. mascula subsp. icarica Tzanoudakis, subsp. nova.

Typus: Insula Icaria, prope pagum Monocampion, Tzan. 221 (typus UPA).

A subspeciebus mascula et russi floribus albis, maioribus differt; a subspecie hellenica foliis subtus valde pilosis segmentis secundariis segmenti centralis primoris in 4-7 foliolis divisis differt.

Ins. Icaria: Mons Atheras, prope pagum Monocampion ca. 600 m. Tzan. 2215 (UPA)” (Tzanoudakis, loc. cit. 1977).

Description: Stem glabrous, 30-60 cm. Lower leaves basically biternate, with 9-21 leaflets, obovate broadly obovate or elliptic and shortly acuminate, twice or less than as long as broad. 1-3 of the leaflets sometimes deeply bifid into similar segments; leaves greyish green above, glaucous beneath, glabrous or pilose; terminal leaflets attenuate into a shortly decurrent petiolule. Flowers 10-13 cm across. Petals 5-7, obovate to obovate-orbicular, white. Filaments purplish red. Carpels 1-5, mostly 3, long-tomentose. Style 8-10 mm long, 2 mm broad; stigmatic area 2 mm broad, circinnate from near base. Fl. April - May. 2n: 20.

Type: "Insula Euboea. Kerasovrysi, in ditione pagi Steni." Tzanoudakis 1210 (holotypus UPA). 2n = 20.

Distribution: Isolated colonies in south-western Peloponnese, Attica, Euboea, Andros (var. hellenica) and Ikaria (var. icarica).

a) var. hellenica

STEREA ELLAS (Central Greece). Prov. 76, Evvia (Euboea): district Karistia, place named Vromeri, 1949, Goulimis 16020 (ATH!); district Halkis, S.E. part ofMt. Dirfis, N.N.E. of the village Steni Dirfios, place called Metohi - Dirfios, 750-800 m., 7. v. 1970, Stamatiadou 8480 (ATH!); E.N.E. of the village Steni Dirfios, place called Kerasovrysi, 700-750 m, 7. v. 1970, Stamatiadou. 8439 (ATH!), 19. iv. 1975, Stamatiadou 18237 (ATH! BM!). Prov. 77, Attiki (Attica): Mt. Parnes, between ravine of Parnes and Malakassa, 10. v. 1970, Goulimis 849 (ATH!); Mt. Parnes. N. side, Abies region, c. 700 m, 12. v. 1976, Stearn (BM!).

PELOPONNISOS (Peloponnese). Prov. 85, Lakonia: district Lakedhemona; Mani, place named Haos, v. 1952, Coulimis 16021 (ATH!); Mons Taygetos, Goulimis 7373 (ATH!). Mt. Taygetos, above the village of Spartia 1200 m. Tzanoudakis (UPA).

KYKLADES (Cyclades): Andhros (Andros), Mt. Kouvara, E. of and above the village Arnas, 600-750 m. 5. iv. 1970, Stamatiadou 7778 (ATH! BM! E!) E.S.E. of the village Arnas on W. slopes of Mt. Kouvara, 600-850 m, 23. iv. 1975, Stamatiadou 18328 (ATH! BM!).

b) var. icarica

NISI TOU EJEOU. Prov. Samos: Ikaria, S.W. of the village Ploumario, on N. foothills of Mt. Atheras, 600-650 m, 29. v. 1970, Stamatiadou 9179 (ATH); W.S.W. of the village Mileopon, on N. foothills of Mt. Atheras, 400-600 m, 6. v. 1976, Stamatiadou 19188 (ATH! BM! E!).

Greek habitats: On schistose substrate, moist and shady places with Pteridium in Abies forest, moist places in thickets Ouercus, Acer, Crataegus, Rubus, Pteridium, open rocky slopes, alt. 400-850 m.

External distribution: Sicily.

Apart from P. clusii of Crete and P. rhodia of Rhodes, the only peony of the Mediterranean area recorded before 1977 in botanical literature as having white flowers was the one in Sicily described by C.B. Presl in 1822 as P. flavescens. Presl visited Sicily in 1817 and collected this on the Nebrodi mountains in the north-east of the island. Since then several collectors have gathered the same plant at various places in the mountains of northern Sicily; Lojacono-Pojero described it as “copiosa nell’alta regione de Faggio e boschi della bassa regione ove é pin rara” with “floribus albidis”. Others have also recorded the flowers as white in a living state. The petals, however, become yellowish on drying, whence Presl’s epithet flavescens. This would appear to be the plant recorded as Paeonia folio subtus incano, flore albo vel paillido C.B.P. in Cupani’s Hortus Catholicus 169 (1696).

During the 20th century collections made in Greece, notably by Pinatzi, Goulimis and Stamatiadou, have revealed the occurrence of white-flowered peonies of the mascula group on the Taygetoy range in Laconia (Peloponnese), in Attica and on the islands of Euboea and Andros. These have been named P. mascula subsp. hellenica by Tzanoudakis. There do not appear to be any characters by which these white-flowered Greek peonies can be separated from the Sicilian one.

Recent as the botanical recognition of these white-flowered peonies has been, their existence was noted in Greek manuscripts of the late 17th and early 18th centuries, according to B. Skoubaras (1967; quoted by Papamichael, 1975:114): "The peony is the mother of all herbs. It grows on high mountains in shelter from the wind, under stones or in places paved with stones. Its leaves reach to the ground. Its flowers are white and its root is like a man's finger."

The differences in habit, leaf, flower colour and follicles separating Paeonia mascula subsp. mascula, subsp. russi and subsp. hellenica are such that, adopting a narrow specific concept as Slapf did, they could be maintained as independent species. No differences of comparable magnitude separate the eastern island population of Ikaria distinguished as subsp. icarica from the mainland and western island populations named subsp. hellenica. In a living state both as wild and cultivated plants they look alike. Some plants from Ikaria have leaflets as sparsely hairy below as those of Andros and Evvia. There is indeed a slight difference in the lower leaves, with hellenica distinguished as having 9-13 leaflets with a length-breadth ratio 1.25-1.70 from icarica distinguished as having 11-21 leaflets with a length-breadth ratio 2.0-2.1, but leaves similar in appearance and the number, shape and hairiness of leaflets can be found in plants from Andros and Ikaria cultivated side by side. Accordingly they are here united in the same subspecies, despite much initial reluctance, and reduced to varietal rank within it as var. hellenica and var. icarica. The large flowers of P.mascula subsp. hellenica with their spreading lightly crinkled pure white petals and red and yellow stamens, arising out of bold foliage, place it among the most beautiful of wild peonies. Big many-flowered clumps growing profusely among rocks on an Andros hillside are one of the most memorable floral spectacles of Greece.

3e. P. mascula subsp. triternata content Figs 33,34

P. daurica Andrews, Bot. Repos. 7: t. 486 (1807); Sims in Curtis's Bot-Mag. 35: t. 1441 (1812); F.C. Stern, Study of Paeonia, 70, t. 6 (1946); Cullen & Heywood in Fl. Europ. 1:243 (1964).

P. triternata Pallas ex DC., Prodr. 1:165 (1824), nomen illegit.

P. corallina var. triternata Boiss., Fl. Orient. 1:97 (1867).

P. corallina subsp. triternata (Boiss.) N. Busch in Kusnezow, Busch & Fomin, Fl. Cauc. Crit. 3.ili:10 (1901).

P. mascula var. triternata (Boiss.) Gürke in Richter, Pl. Europ. 2:400 (1903).

Illustrations: Andrews, op. cit. (1807); Curtis's Bot. Mag. 35: t. 1441 (1812); F.C. Stern, op. cit. (1946).

Protologue; “P. corallina [beta] triternata. Foliola ovato-orbiculata saepe suborbiculala obtusa aut vix acuta. P. triternata Pall. Nov. Act. Petr. X, p. 312. P. Daurica Anders. Bot. Mag. tab. 1441!” (Boissier, loc. cit. 1867).

Description: Stem up to 60 cm, glabrous. Lower leaves biternate, rather thick; leaflets usually 9, very broadly elliptic to broadly ovate-elliptic, obovate or almost orbicular, shortly subacuminate, rounded or truncate at apex, 5-9 cm long, 4-6 cm broad, occasionally with some of the leaflets divided into 1 or 2 smaller leaflets, undulate at margin and somewhat upcurved, deep green above, glaucous and usually glabrous below, the terminal leaflets slightly or distinctly stalked; petiolule 5-20 nun. Flowers 7.5-9.5 cm across. Petals 5-8, obovate-orbicular, rose. Filaments yellow (purplish in Turkey). Carpels 2-4, long-tomentose. Style 8-9 mm; stigmatic area c. 2 mm broad, circinnate from base. Fl. May. 2n: 20.

Type: Crimea (see below).

Subsp. triternata sensu stricto is a native of the Crimea and is there diploid (2:10). It has distinctive foliage, the 9 leaflets being often obovate or almost orbicular, with a rather concave upper surface, undulate margin and rounded to truncate apex. It is well depicted by Lilian Snelling in F.C. Stern's Study t. 6 (1946) as well as in the earlier illustrations of Andrews and Sims, all from cultivated plants. These characters may appear individually but apparently not all together within mascula populations outside the Crimea and the status of such plants, which are covered by the above description, is doubtful. They have more acute leaflets than subsp. triternata senso stricto. They occur on the eastern Aegean islands of Samos and Lesvos, together with subsp. mascula, which is tetraploid (2n: 20). The following gatherings exemplify them:

Lesvos: on the way to the Sanatorium of Ajiassos. 15-20. v. 1952, Goulimis 16032 (ATH!). Samos: Kierki, Forsyth Major (K!); Mt. Kerketefs, 200 m. S and below Profitis Ilias chapel, 2. v. 1968, Stamatiadou 2666 (ATH!).

Some morphological overlap between subsp. triternata and subsp. mascula in Anatolia has led us to give it subspecific rank here. Possibly the Greek material is best designated as "triternata-approaching" (ad subsp. triternatam accedens).

A variant of P. mascula occurring in Romania with typical mascula but approaching triternata in its obtuse or subacute leaflets was named P. corallina var. triternatiformis Nyárády in Savulescu, Fl. Republ. Pop. Romane 2:403, 675, pl. 63 fig. 2 (1953). Population studies are needed to clarify the status of taxa within this mascula complex. If the Crimean peony is accepted as specifically distinct from P. mascula, its nomenclaturally correct name is P. daurica published by Andrews in 1807. No such peony occurs in the region of south-eastern Siberia and adjacent north-eastern Mongolia formerly known as Dauria or Davurica. It was introduced into cultivation in England from the Crimea, the Taurica Chersonesus or Taurica of Ancient Greek and Roman geographers, and the epithet daurica adopted by Andrews is obviously a misrendering of taurica by a gardener. Pallas, who lived in the Crimea from 1795 to 1810, originally named the species P. triternata but never validly published this name; when De Candolle adopted it in 1824, it was antedated by the name P. daurica and made illegitimate. At a varietal and subspecific level, however, the correct epithet is triternata adopted by Boissier in 1867 and Busch in 1901.

4. P. clusii content Plate 13; Figs 7,35

P. cretica Tausch in Flora (Regensburg) 9:88 (1828); non Sabine (1824).

P. peregrina var. glabra Boiss., Fl. Orient. 1:97 (1867).

P. peregrina var. cretica Huth in Engler, Bot. Jahrb. 14:270 (1891).

P. officinalis var. glabra (Boiss.) Hayek, Prodr. Fl. Penins. Balcan. 1:298 (1924).

P. clusii F.C. Stern & Stearn in Curtis's Bot. Mag. 162: t. 9594 (1940); Rechinger f., Fl. Aegaea, 177 (1943); F.C. Stern, Study of Paeonia,

102, t. (1946); Cullen & Heywood in Fl. Europ. 1:243 (1964).

P. clusii subsp. clusii; Tzanoudakis, Cytotaxon. Study Paeonia in Greece, 23 (1977).

Illustrations: Curtis's Bot. Mag. t. 9594 (1940); F.C. Stern, op. cit. (1946).

Protologue: "6. Paeonia cretica: herbacea; germinibus 5 tomentosis; foliis ternatimsectis, segmentis 5-nacontentinnatifidis, laciniis lanceolatis acuminatis basi decurrentibus utrinque glabris.

P. cretica. Clus. hist. 1. p. 281. DeCand. syst. veg. I p. 394.

P. folio subtus incano flore albo v. pallido. C. Bauh. pin. 323.

Habitat in montibus Sphakeoticis Cretae, Sieber.

Steht der P. anomala L. am nächsten. Der Stengel ist aufrecht unbehaart. Die Abschnitte der Blätter 5-zahlig, die Lappen alle ganz lanzettförmig lang zugespitzt ganz unbehaart, unten etwas blasser aber nicht graugrün. Die Blume weiss (Sieber), die Fruchtknoten 5 aufrecht weissfilzig, die Narbe bogenförmig-purpurroth." (Tausch, loc. cit. 1828).

Description: Stem glabrous 20-30 cm, pink or purplish. Lower leaves biternate, leaflets dissected into 30 or more very acute segments which are themselves lobed and toothed, totalling 40-80 leaflets and lobes that are narrowly oblong-lanceolate, acute to acuminate, usually glabrous on both surfaces, firmer textured than in P. rhodia. Flowers 7-10 cm across. Petals 6-8, obovale to obovate-orbicular, white, rarely flushed pink. Filaments pink. Follicles 2-5, shortly white-tomentose; style 7-8 mm, stigmatic area 1.5-2 mm broad, circinnate only in upper ½ - 3/4, Fl. April - May. 2n: 10. (W. Crete and Karpathos), 2n: 20 (E. Crete).

Type: Crete, “in montibus Sphakeoticis Cretae;" Sieber (K! Praha. Holotype!)

Distribution: Endemic to Crete and Karpathos.

Habitats: Pinus halepensis forest, Cupressus sempervirens var. horizontalis forest, Pistacia lentiscus macchie, dry calcareous river beds and among rocks. Altitude 200-190 m. P. rhodia differs from it in leaf shape and texture.

The existence of a white-flowered peony on Crete was recorded in 1553 by the adventurous roving French naturalist Pierre Belon (1517-1564) and confirmed in 1593 by an Italian doctor Onorio Belli (d. 1604), resident in Crete, a correspondent of Clusius as was also a Florentine nobleman Matteo Caccini (1573-1640) who grew various plants from Crete and Constantinople. Letters to him from Clusius were edited and published by P. Ginori Conti in 1939. Thus, it is unlikely that the Paeonia flore albo simplici listed and grown by Caccini could have come from anywhere but Crete. Venetian trading activity made communication between north Italy and Crete easy and frequent. For two centuries thereafter the Cretan peony remained obscure, but in 1817 a Bohemian plant-collector, Franz Wilhelm Sieber (1789-1844), visited Crete and gathered specimens in the White Mountains. His compatriot Ignaz Friedrich Tausch (1793-1848) recognized them as belonging to a new species which he described in 1828 as P. cretica. Unfortunately between 1601 and 1824 the Clusian name P. cretica had become associated with another peony having pink flowers and hairy less divided leaves, long cultivated in Oxford and unknown in Crete, Andersen's P. arietina var. oxoniensis. This was described and illustrated in the Botanical Register 10: t. 819 (1824) as P. cretica Sabine by John Lindley. Hence Tausch's name P. cretica (1828), being a later homonym, could not legitimately be used for the Cretan peony, which accordingly needed a new name. When this came to Stearn's attention in 1935 he renamed it P. clusii and between 1935 and 1939 annotated as such the specimens in the British Museum, Cambridge, Gothenburg, Prague, Upsala and Vienna herbaria, intending himself to publish this manuscript name later. Before then it had been validly published independently by F.C. Stern and K.H. Rechinger.

Few scholars offer a nobler example of devotion to botany, despite ill-health, crippling accidents and poverty, than Charles de l’Escluse (1526-1609), who published in Latin as "Carolus Clusius"; writing in Italian he signed himself "Carlo Clusio". His many-sided curiosity, aided by his grasp of Latin, Creek, Italian, Spanish, Portuguese, German, French and Flemish, took all the period's learning in its survey; he was a master of the art of plant description. Accordingly, on account of his contribution to knowledge of peonies and a vast number of other plants, it seemed fitting to name this peony clusii for him. To their lasting credit the Curatores of the young University of Leiden induced him at the age of 66 to accept the post of director (Praefectus Horti) of the Leiden botanic garden and to spend his last years happily and fruitfully in Holland. After Sieber's visit Paeonia clusii was collected by Raulin, Heldreich and many others and had been introduced before 1920 into English gardens by Aubyn Trevor-Battye, author of Camping in Crete. For many years its specific distinctness went unrecognized, Boissier having named it P. peregrina var. glabra. In 1886 a Scottish doctor interested in the scientific exploration of the Aegean archipelago, Charles I. Forsyth Major (1843-1923), found it on Karpathos and recorded it from there in 1895. Acquaintance with this peony in a living state made evident its well-marked distinctive features.

With its usually cerise-tinged stems, finely cut leaves and large white clove-scented flowers, P. clusii is the most elegant of the Creek peonies.

5. P. rhodia content Plate 14; Figs 36, 37

P. rhodia Stearn in Gard. Chron. Ser. 111. 110:159, Fig. 77 (1941): F.C. Stern, Study of Paeonia 84 (1946); Davis & Cullen in Fl. Turkey 1:205 (1965); Phitos & others. Wild Flowers of Greece 53, fig. 37 (1965); Goulandris, Goulimis & Stearn, Wild Flowers of Greece, 24, t. (1968).

P. clusii subsp. rhodia (Stearn) Tzanoudakis, Cytotaxon. Study Paeonia in Greece, 25 (1977).

Illustrations: Stearn, loc. cit. (1941); Phitos & others loc. cit. (1965); Goulandris, Goulimis & Stearn, loc. cit. (1968).

Protologue: "PAEONIA RHODIA Stearn, species nova. Syn. "P. corallina" sec. Boissier in herb. et apud Buser, Fl. Orient. Suppl. 21 (1888), pro parte quoad plantam insulae Rhodi; non Retzius. Herba perennis, humilis, praeter carpellos glabra. [Radices crassae, cylindricae, sed ut videtur non tuberosae]. Caulis florifer uniflorus, ruber, folios 3 (-4) gerens 25 [-35] cm. altus. Folia plerumque biternata, segmentis haud raro bilobatis, lobis ultimis [7-] 8-22 [-30] plerumque anguste ovatis, lanceolatis vel anguste lanceolatis parvis acutis vel acuminatis basi abrupte vel gradatim contractis saepe non confluentibus, 1.5-10 cm longis, 0.7-3.8 cm latis, petiolis petiolulisque rubris. Flos patens c. 85 cm diametro, fragrans (fide E. Landby). Sepala 4, extimum lanceolatum, intinium rotundatum emucronatum nervo medeano inconspicuo. Petala alba, obovata vel late obovata, c. 3.5-4.5 cm longa, 1.5-2 cm lata. Staminum filamenta rubra, c. 3-7 mm longa; anthera flava. Carpella 2-3, ad anthesin erecta et c. 1.5-2 cm longa [serius patentia et 2-2.5 cm longa], ovariis tomentosis, stigmatibus recurvatis sessilibus rubris. [Planta diploidea, chromosomatibus 10].

Distribution: island of Rhodes, eastern Mediterranean, in mountainous places at about 500-600 m.

Herbarium specimens: Nadelholzwälder am Monte Profitze, c. 600 in [?Profeta Elia near Arcangelo or Prophylia], 1938, Engelhardt et. Landby (herb. Kew typus); collines du Mt. San Elio pres Salakos, 1870, Bourgeau, PI. Rhodes 2 (herb. Kew);

in monte Prophet Elias prope Salakos. in saxosis calc. faucis occidentalibus, c. 500 m, 1935, Rechinger 7226 (herb. Naturhist. Mus. Wien);

sine loc. specif., Crane, cult. hort. Bowles (herb. Kew; lierb. Brit. Mus., London).

With regard to these localities it should be noted that among the Greeks the prophet Elijah (Hagios Elias) is the patron Saint of mountains (cf. Hastings, Dict. Bible, I, 691:1898) and consequently many mountains in Greece and the Aegean islands nowadays bear his name." (Stearn, loc. cit. 1941).

Description: Stem glabrous, reddish, 28-35 cm. Lower leaves biternate. with all or nearly all leaflets divided, totalling 9-29 divisions; terminal leaflet of each trichotomy deeply trifurcate, lateral leaflets usually bifurcate, entire or divided into 3 or 4 lobes. Leaflets and segments ovate to oblong-elliptic or lanceolate, acute to shortly acuminate, thin textured, 2.5-1 cm long, 0.7-3.7 cm broad, green above, paler below, glabrous, subpetiolate or sessile. Flowers c. 7-8 cm across. Petals 6-8, obovate-orbicula, white. Filaments red. Carpels 2-5, tomentose. Style c. 10 mm; stigmatic area 2-2.75 mm broad, strongly circinnate in upper 3/4. Fl. February - April 2 n:10.

Type: (Rhodes) Nadelholzwälder am monte Profitze, c. 6000 m, 1938, Engelhardt & Landby (K!).

Distribution: Endemic to Mount Profitis Elias on the island of Rhodes.

DODEKANISOS (Dodecanese): Rhodes (Rhodes): Collines du Mt. San Elio près Salakos, 1870, Bourgeau, PI. Rhodes 2 (K!); prope Salakos, c. 500 m, 1935, Rechinger (W); Monte Profitze, c. 600 m, 1938, Engelhardt & Landby (K, holotypus); above Archangelos, 350 m, Davis 40312 (R!); Mt. Profitis Elias, E.S.E. slope of summit, 550-700 m. 27 iv. 1969, Stamatiadou 5792 (ATH! BM!).

Habitat: Forest of Cupressus semoervirens var. horizontalis on limestone, rarely in phrygana. Altitude 350-630 m.

This is closely allied to P. clusii of Crete and Karpathos but well distinguished in leaf characters.

Specimens collected in fruit on Rhodes by Emile Bourgeau in 1870 made known the existence of a wild peony on the island but its flowers were unknown until 1938 when Mrs Elsa Landby, a Swedish lady living at Eynsford, Kent, England, sent a pressed flowering specimen to Kew for identification. She had brought back a plant for her garden. Unfortunately she died not long afterwards but her husband kindly permitted Stearn to fix root-tips at Eynsford from which L.F. LaCour determined it as having 2 n:10, hence a diploid; Tzanoudakis has confirmed this number. E.A. Bowles had earlier received a plant from Rhodes, stated to be white-flowered, in 1930; he managed to keep it alive for many years hut it never throve and never flowered again. Convinced of its distinctness Stearn in 1941 described it as a new species.

CHAPTER V CULTIVATION content

The Creek peonies are herbaceous perennials which die down to a rootstock every year but survive the hot summer by means of their fleshy roots. They are easy to grow in a deep well-drained soil provided vigorous stronger-growing plants do not overpower them; at Kifissia seedings of Acanthus mollis tend to invade clumps and endanger them unless weeded out. Wild peonies occur on calcareous formations, often on rocky slopes and in slight shade from bushes and trees; under cultivation they thrive better in limy than acidic soil. All the Greek species are winter-hardy at Kifissia north of Athens; here they do well in partial shade reminiscent of that from rocks and bushes in their native habitats. Further north, as in England, they are best planted out in the open but sheltered from excessive wind; the wall-enclosed herbaceous plant area at the Royal Botanic Gardens, Kew displays large clumps of peony species; here, in England, the two southernmost Greek species, Paeonia clusii and P. rhodia, have proved difficult to maintain. Peonies should be disturbed as little as possible and allowed to spread gradually into big clumps. The most spectacular Balkan species is P. peregrina, with deep red bowl-shaped flowers, well known in European gardens and esteemed for nearly four centuries, having been introduced into Austrian gardens by way of Constantinople (Istanbul) in the last quarter of the 16th century. The old double red variety 'Rubra' of P. officinalis, not a Greek species, is hardy out of doors even in northern Sweden,where it is known as 'Bondpion'; Nils Hylander considered this, which was known to l'Obel in 1581, as a hybrid between P. offlcinalis and P. peregrina (cf. Hylander, 1938) but that is improbable. The white-flowered subspecies hellenica of P. mascula from Andros starts so early into growth in southern England that it is liable to be damaged by frost but otherwise is as easy to grow as the type subspecies of P. mascula and P. officinalis, the 'male' and 'female' peonies of the old herbalists, grown in Central Europe during the Middle Ages. Seeds should be sown as soon as ripe; their oily content probably reduces their longevity. Seedlings take four to five years to reach flowering state. Plants can also be propagated by division, cutting the rather woody rootstock between the dormant buds so that each section has an adequate number of fleshy roots for renewed growth.

CHAPTER VI BIBLIOGRAPHY content

Anderson, F. 1983. German Book Illustration through 1500. Herbals. New York.

Anderson, G. 1818. A monograph of the genus Paeonia. Transactions of Linnean Society of London 12:2-18-290.

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Broussalis, P. 1978. The genus Paeonia in Greece. ^^^^ 14:10-14, 38-39.

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Carniel, K. 1967. Über die Embryobildung in der Gattung Paeonia. Oesterreichische Botanische Zeitschrift 114:4-19.

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Creutzburg, N. 1966. Die südägäische Inselbrücke, Bau und geologische Vergangenheit. Erdkunde 20:20-30.

Cronquist, A. 1981. An integrated System of Classification of Flowering plants, 299-302.New York.

Cullen, J. & Heywood, V.H. 1964a. Notes on the European species uf Paeonia. Feddes Repertorium 69:32-35.

Cullen, J. & Heywood, V.H. 1964b. Paeonia. In Tutin, T.G. & others, Flora Europaea 1:243-244.

Davis, P.H. & Cullen, J. 1965. Paeonia. In Davis, P.H. and others, Flora of Turkey 1:204-206.

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Greuter, W. 1977. Chorological additions to the Greek Flora, 1. Candollen 32:21-49 (Reissued in OPTIMA Leaflets no. 38; 1977).

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Hylander, N. 1938. Om bondpionens uppkomst och vetenskapliga namn. Lustgarden 18-19: 69-76.

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APPENDIX Dioscorides on Paeonia content

(Pedanii Dioscorides De Materia medica, edidit Max Wellmann 2:149-150; 1906).

III. 140. Paeonia or glycyside which some name pentorobon, dactylos idaeos, the root paeonia, others aglaophotida. The stem grows two spans high and has many branches. The male has leaves like walnut, the female much divided leaves like smyrnium. At the content of the stem it produces pods like almonds, in which when opened are found many small red grains like the seeds of pomegranate and in the middle five or six purplish black ones. The root of the male is about the thickness of a finger and a span long, with an astringent taste, white, the root of the female has seven or eight swellings like acorns as in asphodel. The dried root is given to women who have not been cleansed (internally) after childbirth. It promotes menstruation (a dose containing root) the size of an almond being drunk; it lessens abdominal pains when drunk in wine. It helps those who have jaundice and kidney and bladder troubles. Soaked in wine and drunk it scontents diarrhoea. Ten to twelve red grains from the fruit taken in dark rough (dry) wine slop menstrual flow and being eaten they ease stomach pains. Drunk and eaten by children they remove the beginnings of stone. The black seeds are good against nightmares, hysteria and pains of the womb when up to fifteen are drunk in mead or wine. It grows on high mountains and foothills.

140 RV. Male paeonia or glycyside, some name pentorobon, orobadion, orobax, haemagogon, pasidee, menogeneion, menion, paionion, Panos cerata, Idaeos dactylos, aglaophotida, theodoreton, selenion, selenogogon of the prophets, phthisis, the Romans casta. Female Paeonia also called aglaophotida. (New translation).

APPENDIX Pliny on Paeonia content

X. Vetustissima inventu paeoina est. nomenque auctoris retinet. quam quidum pentorobon appellant. alii glycysidem. nam haec quoque difficultas est quod eadem aliter alibi nuncupatur. nascitur opacis montibus caule inter lolia digitorum quattuor ferente in cacumine veluti Graecas nuces quattuor aut quinque. inest his semem eopiosum. rubrum nigrumque. haec medetur et Faunorum in quiete ludibriis. praecipiunt eruere noctu, quoniam si picus Martius videat tuendo in oculos impetum faciat.

X. The first plant to be discoveres was the peony, which still retains the name of the discoverer; it is called by some pentorobon. by others glycyside. for an added difficulty in botany is the variety of names given to the same plant in different districts. It grows on shaded mountains, having a stem among the leaves about four fongers high, which bears on its content four or five growths like almonds, in them being a large amount of seed. red and black. This plant also prevents the mocking delusions that the Fauns bring on us in our sleep. They recommend us to uproot it at night-time, because the woodpecker of Mars, should be see the act. will attack the eves in its defence.

[Pliny, Natural History (Loeh Classical Library) 7:156-157 (1955) edited and translated by H.S. Jones).

The woodpecker superstition mentioned here by Pliny and earlier by Theophrastos (page l6) is in some way connected with the ancient pre-christian veneration of the woodpecker as an oracular bird of the God Mars and guardian of sacred groves, presumably the great red-capped black woodpecker (Dyocopus martius syn. Picus martius L.); see E.A. Armstrong. The Folklore of Birds, an Enquiry into the Origin of some Magical Religious Traditions 93-112 (1958).

ACKNOWLEDGMENTS

Grateful acknowledgment is made to the authorities of the British Museum (Natural History), South Kensington, London and the Royal Botanic Gardens, Kew for facilities of studying the material in their charge as also for the provision of photographs of illustrations in early botanic works. The SEM micrographs of pollen grains (Fig. l9, 20) have been made by Mrs. Madeleine Harley (Kew) and Mr. Graham L. A. Heath (B.M.(N.H.)) whose co-operation is much appreciated. To Mrs. Diana Dopheide we are indebted for unstinted help in preparation of the book. The frontispiece depicting the earliest known illustrations of Paeonia come from the Codex Patavinus of Dioscorides in the Seminario Vescovile, Padua through the gracious co-operation of the director. The maps have been prepared by Miss Kiki Dima, and Miss Matina Theodorou. Mrs. Ruth Stearn has compiled the index. The format of the book and the task of seeing it through the press have been the responsibility of Mrs. Vasso Zambeli - Koussoula, to whom we are also grateful for many line - drawings. The book owes its origin to Mrs. Niki Goulandris who many years ago urged W.T. Stearn to resume study of the genus Paeonia using the collection of living plants from many Greek localities assembled in the garden of the Goulandris Natural History Museum at Kifissia as well as the continuously enriched material of its herbarium.