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THE GARDENERS' CHRONICLE. November 1, 1941 pp. 158-160

PLANTS NEW AND NOTEWORTHY.

PAEONIA RHODIA: THE WILD PAEONY OF RHODES.

William T. Stearn

ALTHOUGH for nearly two thousand years the island of Rhodes, in the eastern Mediterranean, has been under alien rulers —Roman, Saracen, Knight Hospitaler and Turk, Italian—many centuries before the birth of Christ it became an important centre of Greek civilization, and Greek-speaking its people have remained. In ancient times, according to the Greek grammarian, Hesychios of Alexandria (quoted by Cecii Torr, Rhodes in Ancient Times, 75: 1885), they worshipped Zeus in the form of Paeon the healer, from whom the genus Paeonia takes its name. There was, however, no evidence of any Paeony growing upon the island until 1870, when a French plant-collector, Emile Bourgeau, brought away specimens in fruit. These came from Mount Hagios Elias, a mountain of about 798 metres on the north-west side of Rhodes, near Salakos. Boissier identified them as belonging to Paeonia corallina (see Boissier, Fl. Orient. Suppl., ed. Buser, 21 pp. : 1888) [The "forma leiocarpa" of Paeonia corallina recorded by Boissier from Kesrouan (Feitroun) can bardly be other tban P. kesrouanensis (Thieb.), S. Thiebaut (1936), syn. P. corallina var. kesrouanensis, Thiebaut (1934), a pink-flowered species with glabrous carpels. Kesrouan is about 25 kilometres north-west of Beirut.] and they were distributed to various herbaria as such. About 1930 an American friend of Mr. E. A. Bowles, Dr. Frederick D. Crane, found a white-flowered Paeony on the island, and sent a living plant to Mr. Bowles, who has managed to keep it alive at Myddelton House, but not to make it thrive and flower again. In Mr. Bowles's garden it starts very early into growth, and is very liable to suffer damage during the winter. Rhodes has a mild climate. In 1935 an Austrian botanist, Dr. K. H. Rechinger, Jun., found a species of Paeony growing at several places among the mountains of Rhodes ; the petals had fallen, but he collected a specimen near Salakos, Bourgeau's locality, which is now in the Natural History Museum of Vienna. Near Arcangelo, on the east coast of Rhodes, he gathered seeds, which he kindly gave me. Plants raised from these seeds still live, but have not proved at all vigorous. The leaves in all these gatherings are completely glabrous. They do not match satisfactorily the leaves of any other Paeony. The one that comes nearest is P. arietina, concerning which it may be well to add a few words.

In herbaria and floristic works the Paeonies that inhabit the mountainous regions of the Near and Middle East are usually placed under two species, P. officinalis (syn. P. feminea, the Female Paeony of the old herbalists) [+ The terms "male" and "female" as applied to plants by the old herbalists are purely metaphorical. When they wished to distinguish two species rather similar in appearance they called the one "male," the other "female," purely for convemence. They did not regard them as a functionally male and female ; mdeed, the existence of sex in plants was hardly realized betöre the seventeenth Century. Paeonia mascula having undivided leaflets, is a more robust-looking species than P. officinalis (P. feminea) with its more divided leaves Many instances of this primitive nomenclature, which has come down from Greek and Roman writers, are given by J. B. Saint-Lager in Ann. Bot. Soc. Lyon XI (1883) 1-48 (1884), and T. A. Sprague, in Journ. Linn. Soc. Bot. XLVIII 87 (1928)].and P. corallina (i.e. P. mascula, the herbalist's Male Paeony). These were the only two Paeonies distinguished by European herbalists down to the sixteenth century. Both have purplish-red flowers. In 1818 George Anderson published an intermediate species as P. arietina, the name (from arietinus, of, or resembling, a ram) referring to its large, recurving mature carpels. Anderson knew it only from garden plants whose history had been lost. Of its occurrence in a wild state he knew nothing. Later botanists have mostly ignored its existence. To the late Dr. Otto Stapf belongs the credit of recognizing wild specimens from Greece and Turkey, mostly labelled P. officinalis or P. pubens, as members of the same species as Anderson's garden-grown P. arietina, and of keeping them apart from P. officinalis. P. arietina thus stands as an independent species widely distributed in Asia Minor; in the Balkans it replaces P. officinalis east and south of Albania, and it may occur in northern Italy. To discuss its variation is not possible here. In general it is a robust-growing plant, much like P. officinalis, with purplish-red flowers and leaves distinctly hairy and glaucous below, and less divided than those of P. officinalis. There are several forms in cultivation, including a white-flowered one which has grown for many years in Sir Michael Foster's old garden at Shelford, near Cambridge; the finest is one with rich amaranth flowers, introduced by Mr. G. P. Baker from north-eastern Turkey. It is a tetraploid species, having twenty chromosomes in its vegetative cells (fide H. N. Barber in litt.).

It was evident from the specimens mentioned above that the Rhodes Paeony did not agree with P. mascula, P. clusii and P. arietina, but a specimen of the plant in flower was lacking until 1938, when a Swedish lady, the late Mrs. Elsa Landby, presented to the Kew Herbarium specimens gathered in Rhodes on "Mte. Profitze." Search through the available maps of Rhodes, notably the very detailed twenty-one sheet Italian Carta dell' Isola de Roai (Florence, 1928), has failed to reveal any place or feature thus named, although the name is certainly a corruption of a Greek or Italian one. Mr. T. Landby tells me that his wife found the Paeony on one of the highest mountains of the island, but he does not remember the exact place. The white petals of Mrs. Landby's specimens confirmed the plant's distinctness. Mrs. Landby uprooted a plant for her garden at Eynsford, Kent, where I have seen it growing. She placed it in the dappled shade of trees so as to imitate its original home ; it has kept alive, but has neither thriven nor flowered again. From root-tips of this plant, gathered by me at Eynsford, and fixed in a 3 : l absolute alcohol—glacial acetic acid mixture, Mr. L. F. LaCour, of the John Innes Horticultural Institution, has determined it to be a diploid (with 2 n = 10), thereby giving a clue to its lack of vigour and further distinguishing it from the robust tetraploid P. arietina (with 2 n == 20). Thus in chromosome-number, as in lack of vigour, general appearance, root-character, flower-colour, glabrousness and variation in leaf-form, this Rhodes Paeony is clearly distinct from P. arietina.

A very remarkable feature of the Rhodes Paeony is its Variation in leaf-foim. (Fig 75). lhaveseen leaves with 7, 9, 11, 13, 14, 16, 17, 18, 22, 24 and 30 ultimate lobes. Bilobing of the lateral segments, sometimes of the terminal Segments, is not infrequent. AU the specimens seen have been glabrous. In a living state the leaves of cultivated plants are somewhat yellowish-green above; they are a lighter greyish-green below but, like those of the Cretan Paeony (P. Clusii syn. P. cretica, Tausch non Sabine), they possess no glaucous bloom removable on handling; they are not distinctly glaucous and hairy below as are those of typical P. arietina. The roots like those of P. Clusii, have been cylindric in all the living plants examined (Crane's, Rechinger's and Landby's) and do not appear to make fusiform tubers. The flowers are described bv Mrs. Landby as scented, whereas those of p arietina, in its white as in its red forms, have the fusty smell characteristic of many Paeonies.

Its near neighbour, P. Clusii of Crete and Scarpanto (Karpathos), is distinctiy clove-scented. This Paeony has likewise a white flower and low growth, but may be readily distinguished by its more finely divided leaves with from eleven (on an uppermost leaf) to seventy ultimate lobes usually decurrent and confluent at base. It is figured in The Gardeners' Chronide, 3rd ser. LXXXIX, 415 (1931 XCIII, 61 (1933) ; Journ. Roy. Hort. Soc LXI, t. 101 (1936) ; and Bot. Mag., CLXII, t. 9594 (1940). It appears to have been cultivated at Florence by Matteo Caccinii, about 1600. The plants in British gardens today come from introductions by Messrs. A. Trevor-Battye G. P. Baker, F. Jekyll, Peter Davis, and Dr A. Q. Wells.

The Rhodes Paeony is not known from outside Rhodes, but may occur in the mountains of adjacent Asia Minor, and perhaps of eastern Crete. Its known range is thus very limited, for the island of Rhodes itself is only forty-five miles long and twenty-two miles broad, .and the Paeony grows only in high places. The mountains of Rhodes, like those of Scarpanto and Crete, are of limestone. The Mediterranean Paeonies appear to be found almost invariably on calcareous formations, and mostly in light woodland or among bushes, scrub and rocks, which screen them from scorching sun. Hence the destruction of forests on Crete and Rhodes has probably led to their disappearance in many places. A French naturalist, Pierre Belon (1517-64), who visited Crete between 1546 and 1549, recorded in 1553 that a white-flowered Paeony grew in humid (presumably shady) valleys near Mount Ida. Today the white-flowered Cretan Paeony (P. Clusii) is only known in Crete from the White Mountains of western Crete and the Lassithi Mountains of eastem Crete, groups about ninety miles apart (Fig. 76) ; no modern collector has found it on or near Mount Ida. It occurs on the near-by island of Scarpanto, whence it was first recorded by Forsyth-Major, in 1895, and where Rechinger has collected it recently. No Paeony is known to grow on the rather flat and barren treeless island of Kasos (Caso), which lies between Crete and Scarpanto. It has been suggested that the Rhodes Paeony might be identical with Thiebaut's P. corallina var. orientalis (Bull. Soc. Bot. France, LXXXI, 114: 1934; Thiebaut, Fl. Libano-Syr., I, 37: 1936), but judging from the type-specimens this Stands between P. corallina and P. arietina, and is certainly distinct from the Rhodes Paeony: it has rose flowers and leaves inconspicuously hairy beneath, and comes from Lebanon.

The Rhodes Paeony is thus an unnamed species, apparently a Rhodian endemic, for which the name of Paeonia rhodia is here proposed. A number of Rhodian coins minted between 408 B.C. and 43 B.C. depict the Rose (see Anson, Numismata Graeca, III, 136-144 : 1912), and the .name of the island may be derived from posov, a Rose. Paeonia rhodia (Fig. 77) has unfortunately little or no garden value, despite its attractive appearance, because it is too difficult to grow. Nevertheless its botanical interest is high.

P. officinalis, a tetraploid species of the Alps and west Balkan Peninsula, has its diploid counterpart in the Cretan species, P. Clusii. P. rhodia is the diploid counterpart of the tetraploid P. arietina. The contrast in area between allied diploid and tetraploid species of Paeonia is paralleled in Tulipa, Tradescantia and other genera. For discussions of this matter and of the taxonomic treatment of closely related diploids and polypioids, see E. Anderson & K. Sax in Bot. Gaz., XCVII, 463-468 (1936), A. Müntzing in Hereditas, XXI, 288-326 (1936). J. A. Nannfeldt. in Svensk Bot. Tidsk., XXXII. 306-312 (1938), and H. A. Senn, in Bibl. Genetica, XII, 298-363 (1938).

The following is its formal description :—

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. Folio 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. 8.5 cm. diametro, fragrans (fide E. Landby). Sepala 4, extimum lanceolatum, intimum rotundatum emucronatum nervo mediano 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 diptoidea, chromosomatibus 10].

Herbaceous perennial, glabrous except for the carpels. Roots moderately thick, cylindric, but apparently not swollen into fusiform tubers. Flowering stem one-flowered, reddish, carrying three or four leaves, 25-35 cm. high. Leaves to 25 cm. long, varying much in form and divided into seven to thirty ultimate lobes (or leaflets) ; lower leaves sometimes biternate with nine entire leaflets, sometimes with the terminal lateral segment bilobed, making ten or eleven leaflets; sometimes ternate with the three main divisions pinnate and the terminal and lower lateral segments of the lateral divisions bilobed, sometimes further divided, making up to thirty leaflets; leaflets shortly petiolulate or almost sessile, narrowly ovate, lanceolate or narrowly elliptic, the base cuneate and rarely confluent with that of another leaflet, 1.5-10 cm. long, 0.7-3.8 cm. broad ; petioles and petiolules reddish. Flower about 8.5 cm. across, scented (according to Mrs. Landby). Sepals four, the outer lanceolate, the innermost rounded with the mid-rib inconspicuous and not reaching to the apex. Petals white, obovate or broadly obovate, 3.5-4.5 cm. long, 1.5-2 cm. broad. Stamen-filaments red, 3-7 mm. long ; anthers yellow. Carpels two or three, at anthesis erect and about 1.5-2 cm. long, later spreading and 2-2.5 cm. long, with the ovaries tomentose, the stigmas sessile, large, red and recurved. Plant diploid (2n == 10).

The details in square brackets in the Latin description are from specimens other than the type-specimen, although the chromosome-number has been obtained from a living plant collected on Rhodes at the same time as the type-specimen.

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

Herbarium Specimens:—

Nadelholzwälder am Monte Profitze, c. 600 m. [? Profeta Elia near Arcangelo, or Prophylia], 1938, Engelhardt et Landby (herb. Kew; typus) ;

collines du Mt. San Eilio 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; herb. 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, Did. Bible, I, 691 : 1898), and consequently many mountains in Greece and the Aegean islands nowadays bear his name.—William T. Stearn.