The Caucasian Representatives of the Genus Paeonia L.
L.M. Kemularia-Nathadse, Trudy Tiflis. Botan. Sada 1961
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Chapter IV
Systematic analysis of Caucasian species of the genus Paeonia L.
Linneus, Syst. pl. (1735) I; Gen. pl. (1737) I; Sp. pl. (1753) 530; De Candolle, Prodr. I (1824) 64; Prantl in Engler et Prantl. Pflanzenfam. III, 2, (1888) 54; N. Busch. in Fl. cauc. crit. Ill, 3 (1901-1903) 7; Grossheim, Fl. Caucasus II (1930) 29; IV (1950) II, Schipczinsky in Fl. USSR, VII (1937) 23; Kolakovsky Fl. Abch. II (1939) 120; Stern. Gen. Paeonia, the Royal Horticult. Soc. I (1946), I, Kemularia-Natadze in Fl. Georgia IV (1948) 4; Rzazade in Fl. Azerbaydjan IV (1953) 29.
Blossoms solitary, actinomorphous, bisexual with a geminate perianth. The calyx lobes are green, leaf-shaped, noticeably unequal, lasting for long at fruits. The petals are large, broader than sepals (calyx lobes), 5-8 in number and in rare cases more than this, purple, pink or yellow, yellowish-whitish. The stamens are numerous, fastened to the disk above a receptacle, spyral-shaped, loose. Pistils are loose, 2-5 in number. The ovary is superior, slightly hairy or bare. The stigmas are laminate, broad, fleshy, coloured. The fruits are follicle-shaped, fleshy and, after getting ripe, they have robust woody walls. The seeds have a fleshy, appendage. The ovule external envelope is longer than the internal one. The plants are large, perennial, with alternate double-tripple ternately or pinnately parted leaves.
Typus: P. officinalis L. a, mascula et b. femina.
According to the newest monograph by Stern (1948), the genus Paeonia comprises about 45 species, propagating mostly in the mountain regions of the East Asia, South Europe, Caucasus, Japan and West states of the North America, of which 13 species are typical for Caucasus, in accordance with the data availed by us. All representatives of the genus are high ornamental.
Table for defining Caucasian species of the genus Paeonia L.
1. Blossoms yellow, cream, yellowish white (yellow after being dried out); ovary and fruits are naked or tomentose-downy; leaf-lobes are always-broad, smooth-edged ....... 2
- Blossoms red, purple, pink; ovary and fruits are always tomentose-downy. The leaf-lobes are broad, smooth-edged or narrow, linear-lance-shaped, linear or almost filiform, incised or dissected at the edges .......... 6
2. Ovary and fruits are bare, stigmas are tapered into more or less long styles ....... 3
- Ovary and fruits are tomentose downy; stigmas are sessile or upon short styles ......... 4
3. Leaves at both sides green, often lustrous from above, bare or grayish-green from beneath and sparse-hairy; their leaf lobes are large, broad obovate or oval, long-tapered at the base. The blossoms are broad-patent, with pale-yellow, almost white petals ............ 1. P. macrophylla (N. Alb) Lomak.
- Leaves at the upper side dull-green, bare, at the inferior side they are grayish, densely downy; the leaf-lobes are ovate, rounded or nearly heart-shaped at the base. The blossoms are not wide-patent, their petals are yellow ............ 2. P. Steveniana Kem.-Nath.
4. Leaves at both sides glaucescent, their lobes are obovate, long-tapered at the base, rounded or bluntly short-pointed at the apex, bare or with nearly unnoticeable short puny hairs at the lower side. The stigmas are pink; ovary is covered with yellowish curling hairs; filaments are yellowish-green; petals are sharply concave, yellow; blossoms are not wide patent .................. 5. P. Mlokosewitschii Lomak.
- Leaves at both sides green, or blue-greyish at the lower side, naked or covered with long hair at the lower side, filaments are flesh-coloured, purple .......... 5
5. Leaves thin, green at both sides, shining, bare or downy at the lower side; leaf-lobes are oval or obovate, long-tapered at the base. The petals are whitish-yellowish, less frequently pinkish-yellowish, oval with a broad claw 3. P. tomentosa (Lomak) N. Busch
6. Leaves parted into broad, ovate, oval lobes (leaflets)
- Leaf lobes are narrow, lance-shaped, narrow-lance-shaped, linear or almost filiform
7. The ovary and fruits are with pink or red tomentose pubescence, stigmas are pink, blossoms are pink, leaves are grey-blue at both sides, leaf-lobes' are rounded, not large ............ 9. P. tritenata Pall.
- Ovary's and fruits pubescence is never pink or red, stigmas are violet-red or flesh-coloured ........ 8
8. At both sides the leaves are green, lustrous and after having been dried out, they become almost membranous-leather-like. Their lobes are broad-oval or broad-ovate, or nearly rhomb-shaped, being abruptly short-narrowed at both sides. The blossoms are not wide-patent (open), with pink or pinkish-reddish, obovate, sharply concave petals ....... 6. Ruprechtiana Kern. -Nath.
- At both sides or from below, the leaves are glaucescent, their, lobes are oblong-ovate, oblong-elliptical, obovate, tapered or rounded at the base ........... 9
9. Leaves glaucescent at both sides, with smooth lobes having slightly protruding ribs, oblong-elliptical. The blossoms are pink, wide patent (open), with obovate slightly concave petals ....... 8. P. lagodechiana Kem.-Nath.
- From beneath the leaves are glaucescent, rough with sharply protruding veins. The blossoms are purple, reddish-pink, wide-patent, with oval or obovate sharply concave petals ........ 7. P. caucasica N. Schipcz.
10. Blossoms pink or reddish-pink, large, leaves are thrice pinnately incised or dissected into narrow lanceolate lobes. It grows upon the margins of sun-lit forests ............ 10. P. Majko Ketzch.
- Blossoms purple or bright-red, leaves are manifold dissected into narrow-linear or nearly filiform leaflets 11
11. Ovary and fruits with red tomentose pubescence. The blossoms are dark-purple with obovate petals. The leaflobes are very narrow, 1-2 mm wide, long, nearly filiform. The radical thickening are oblong, upon short pedicels
11. P. tenuifolia L.
- The ovaries and fruits are with yellowish or whitish-grayish tomentose pubescense, leaf-lobes are wider, narrow-linear, 3-5mm wide. The radical thickenings are oval, rounded, upon long pedicels 12
12. Leaves grayish, covered along veins at the upper side with sharply protruding rows of hairs. The blossoms are bright-red 13. P. Biebersteiniana Rupr.
- Leaves green, naked. The blossoms are dark-purple 12. P. carthalinica N. Ketzch.
Sect. Flavonia Kem.-Nath. h. I. comb. nova Syn. Sect. Paeon DC. Prodr. (1824) 65 p.p. Sect. Albiflorae Salm. - Dyck. Hort. Dych (1834) 365, 366 pp. Sect. Palearcti-ceae. herbacea Huth. in Engler Bot. Jarhb. 14 (1892) 265 pp. Subsect. Foliolatae Stern. A Study of the Gen. Paeonia in Royal Horticult. Soc. (1946) 2. Flores lutei. ochroleusi. Ovarium fructisque glabri vel tomentosi. Folia bivel-triternata foliolis latis integris. Plants perennials Caucasi et Asiae orientalis incola. The blossoms are yellow, cream, get yellow in result of drying out. The ovary and fruits are naked or tomentose hairy. The leaves are thrice ternate with broad smooth-edged lobes. The plants are perennials growing in Caucasus and the East Asia. Type: P. macrophylla (N. Alb.) Lomak. and P. Wittmanniana Hartwiss.
Series Macrophyllae Kem.-Nath.
Syn. Series Corallinae Kom. in Fl. USSR VII (1937) 28 pp. The leaf-lobes are very broad, ovaries and fruits are naked. Foliola lastissima. The type P. macrophylla (N. Alb.) Lomak.
1. Paeonia macrophylla (N. Alb.) Lomak.
Lomakin in "Trudi of Tifl. Bot. Sad" II (1897) 282, Lipsky "Fl. Cauc." (1899) 213; Komarov and Schipsky in "Fl. USSR" VII (1937) 30.
Syn. P. corallina. var. Wittmanniana forma Macrophylla N. Alb. Prodr. Fl. Colchid. (1895) 15; P. Wittmanniana Stev., forma macrophylla (N. Alb.) N. Bush in Fl. Cauc. critica III, 3 (1901-1903) 13; P. Wittmanniana Hartwis ex Lindl. var. macrophylla (N. Alb.) N. Bush ex Grossheim, "Fl. Cauc." 2 (1930) 91, F. C. Stern in Journ. Royal. Hort. Soc. 68 (1943) 126; L. Kemularia-Natadze in "Fl. Gruzia" IV (1948) 6; Grossheim, Opred. (1949) 44, Fl. Cauc. IV (1950) 13; Stern, A Study of the Gen. Paeonia in Royal Hortic. Soc. (1946) 59; P. Wittmanniana, Kolakovsky "Fl. Abch." II (1939) 121 pp.
Perennials. The rhizomes are nearly horizontal, branchy with fusiform radical thickenings; stems are long, 100-150cm high, firm, branchy; leaves are green at both sides or grayish from beneath, naked or downy from beneath. The leaf-lobes are large, 12-15cm long, obovate, oblong-oval, tapered at the base and apex or long acuminate at the apex only. The blossoms are wide patent, with cream, yellowish-white slightly concave petals; anthers and filaments are yellow; or filaments are dark-violet at the base; ovary is naked, long, narrow, ovate-cylindrical, gradually tapering into a long style at the apex; stigmas are dark-violet, one-sidedly laminate (very often the whole pistil is coloured in violet). The fruits are large, long, oblong, spreading, after getting ripe they are pendent. The seeds are bluish-black, (V-VI).
Habitat. From the upper mountain belt up to the subalpine belt, in forest and on margins. Original area. Georgia, Adjaria. A slope of mountain Chkvish-Tavl. the village Agara.
Type. It has been described from live samples grown in the late Caucasian department of the Tiflis Botanical Garden from seeds, procured by N. Albov.
Studied samples. Georgia, Abkhazia. The outskirts of Gagri, Schavrov!
Guria, the Adjar-Imerinsky ridge, the Guriysky ridge. The outskirts of Bakhmaro, 6/VII-1914, E. Kikodze!
The Adjar-Imeretinsky ridge, the border between shrubbery thickets and alpine meadows by Zotis-Keli, 7/V-1914, E. Kikodze!
Adjaria, mountain slope of Chakvis-tavi near the village Agara, 14/VIII-1893, Albov!
Cultivated at the Cauc. dpt. of Tifl. BG, 9/IV-1902, Fomin!
The Adjaro-Guriysky ridge, by the pass up to Gomis-mta, a meadow in the midst of subalpine forest, at the west slope, 13/VIII-1956, A. Dolukhanov and A. Kazarova!
Geographical type: Mountain Kolkhidsky. Main distribution: West Transcaucasus.
Remarks. This species was described in 1895 by N. Albov and named P. corallina var. Wittmanniana forma macrophylla N. Alb. Lomakin was quite right, having separated it from the red-flowered peonies and determined it as a self-dependant species, different from P. corallina and P. Wittmanniana Stev.
Afterwards, with exception of Lipsky (1897) and authors of "Flora USSR" (1937), botanists considered this plant a variety or form of the species P. Wittmanniana, accepting a point of view of Steven, or Hartwiss, or Lindley. Through the magazine "Flora Gruzii" (1948), we expressed the opinion of the majority of botanists, considering P. Wittmanniana Stev. and P. Wittmanniana Hartwiss ex Lindley, as one species. But at present, after it has been found out that the name P. Wittmanniana (see references about P. Wittmanniana Hartwiss) denotes two different species and the characters of the Adjarian peony are transitional between the characters of the two above-mentioned peonies,.we were compelled to discard our previous point of view and recognize the Adjarian peony, as a self-dependent species.
Undoubtedly, P. macrophylla (N. Alb.) Lomak. with its naked fruits and peculiar form of ovary is much akin to the species of Steven (from our point of view). Nevertheless, the former species distinctly differs from the latter with its form, broad-patent blossoms, petals colour (more pale) and lobes form at both sides of nearly mono-coloured green leaves (in most typical cases).
Both species differ from each other with areas of distribution, just for instance, P. macrophylla (N. Alb.) Lomak. may be met in Abchazia also, where P. Wittmanniana Hartwiss grows, so that it does not abandon the region of Kolchida (its area may be Lasistan), and as to P. Steveniana Kem.-Nath. (P. Wittmanniana Stev. non Hartwiss), it grows in forests of the East Georgia, the regions adjoining the West Transcaucasus, which are much influenced by the Kolchidsky-floristical climatic conditions (Mesketia, Borjom-Bakuriani and the South-Osetia).
It should be emphasized that the downy-fruited peonies grow separately from the bare-fruited peonies (Lagodech, Talish for the former peonies; Adjaria, Guria, Imeretia, Megrelia, Meschetia, Kartly, the South Osetia for the latter peonies) and these two peonies are never met growing together in other regions of Caucasus.
2. Paeonia Steveniana (Stev.) Kem.-Nath. nomen novum.
Syn. P. Wittmanniana Stev. in Bull. Soc. Nat. Mosq. ser. Ill (1848) 275; Boiss. Fl. Or. I (1867) 97 pp. ; Rupr., Fl. Cauc. (1869) 46; N. Busch in Fl. Cauc. crit. Ill, 3 (1901-1903) 12; Grossg. Fl. Cauc. II (1930) 91; Schipch. in Fl. USSR, VII (1937) 29; Kemularia-Nathadse in Fl. Gruz. IV (1948) 5, P. Wittmanniana var. nudicarpa Schipcz. in Not. Syst. Herb. Hort. Bot. Petrog. 2 (1921) 44; Stem, Monogr. in Jour. Horticult. Soc. I. (1946) 59; Grossg. Fl. Cauc. (1950) 12. XVI.
The plants are large, 50-100cm high or somewhat higher; rhizomes are branchy with dark-brown, long spindle-like root thickenings; the stem is firm, cylindrical or slightly ribbed; the leaves are double thrice ternate, green, bare at the upper side, grayish, with more or less dense long white hairs at the lower side; leaf-lobes are ovate or oblong-oval, of which the lateral lobes are with a rounded or nearly heart-shaped base, upon short petiols or being nearly sessile, narrowed at the apex; the terminal lobe is large and, usually, tapered at both tips. The blossoms are not broad-patent, with sharply concave petals, filaments are yellow as a whole, or flesh-coloured as a whole or up to the middle; ovary is oblong-ovate, naked; stigmas are flesh-coloured, tapered, with a more or less long style. The fruits are naked, oblong-ovate or oval, curvedly spreading.
Habitat. From the upper montane belt to the subalpine belt, in forests, on margins, in shrubberies. Original habitat: Georgia, Meskhetia, between Akhaltsikh and Atskhur.
Type. "In the shady gorge between Kartalinia and Akhaltsikh, not far from Atskhur. V, 1840, Wittmann". The description was obtained from live samples brought up in the late Caucasian dept. of the Tiflis Botanical garden.
Topotype. Atskhur. Akhalts. 25, VI, 1903, A. Fomin. It is kept in Herbarium of the Institute of Botany of the Georgian Academy, Tbilisi.
Studied samples. Georgia, Abchazia, the outskirts of Gagri, Schavrov!
Svanetia, the southern slope of the Dadianovsky ridge, Akinfiev!
Megrelia. The ridge Okhachkuye, lime-stones, 9. VII. 1956, Kemularia-Nathadse, Kutateladze, Mandenova, Mikeladze, Dolidze! Djvari, 1893! Albov, 1956! Khintidze and Mikeladze!
Mnt. Migaria, 22.VI.1957, Khintibidze and Mikeladze! E. and M. Sokhadze!
Mnt. Askhi, 19. V. 1957, Kemularia-Nathadse, Khintibidze and Mikeladze.!
Racha-Lenchumi, the Khvamli 13.VI.1958 Kemularia Nathadse, Khintibidze and Mikeladze! 1957. E. and M. Sokhadze!
Nakeralsky ridge 26.VI.1928 Kemularia-Nathadse, G. Chichinadze!
Imeretia, the Nakeralsky ridge, near Chischkara 24.VI.1928 Kemularia-Nathadse. G. Chichinadze and V. Djindjikhadze!
Nakeralsky ridge 1951, A. Kutateladze!
Kartly. Borjom, "Kupalische", in forest, 15.V. 1913, Sosnovsky!
Outskirts of Borjom, the Baniskhevsk gorge, 27. V. 1920!
the ridge Dashiris-seri, the north slope, 8.VI.1920, Kozlovsky!
Bakuriani, 6.VI.1928, Mukheri! 10.VI.1913, Kozlovsky; Sakochavi! 25. V. 1888, Medvedev! 17.VIII.1946, V. Nakhutsishvili!
in forest by the Sakochavsky lakes. II.VI.1913. Kozlovsky! 17.VII.1917. L. Kemularia! VIII. 1935, I. Barnabishvili!
South Ossetia. Mut Sirkh-Laberta, a forest in the direction of Hohenacker, 2.000m, 21.VIII.1928! Khodze-Khor, 19.VII.1930 E. and N. Busch!
Meskhetia. B. Ankur. "The shady gorge", V.1840, Wittmann!
Atskur, 25.VI.1903, A. Fomin!
Geographic type. Montane Kolchidsky. General distribution. Endemic.
Remarks. This species was given an improper name, unfortunately (see P. Wittmanniana Hartwiss). It was described by Steven from the samples collected by Wittmann in "the shady forest between Kartalinia and Akhaltsikh near Atskhur" in 1848. But earlier, in 1846, the identical name was given to a peony from Abchazia which was described by Hartwiss and Lindley. In this case, the priority of authorship and name belongs to Hartwiss, and the Steven's peony is to be given another name. As Steven was first who paid attention for this plant and described it as a new species, we name it in honour of the academician Steven. This species differs from the P. Wittmanniana Hartwiss ex Lindley not only with bare fruits, but also with the form of ovaries and stigmas, with leaves which are grayish and densely downy at the lower side and which lobes are sharpened at the apex and often nearly rounded at the base. It resembles P. macrophylla (Alb.) Lomak. with the absence of pubescence upon ovaries and fruits and with their form. With its leaflobes form, it has some likeness with the P. tomentosa (Lomak.) N. Busch from Talish from which it distinctly differs with the naked fruits, petals form and colour and separated area of distribution, nevertheless.
Series Wittmanniana (Stern) Kem.-Nath. Wittmanniana group Stern A study of the Gen. Paeonia the Royal Horticult. Societ. (1946) 53. Series Obovatae Kom. in "PI. USSR" VII (1937) 26 pp. Ovaries and fruits are tomentose-downy. Ovarium fructusque tomentosi. Type. P. Wittmanniana Hartwiss ex Lindley
3. Paeonia Wittmanniana Hartwiss
Hartwiss ex Lindley Bot. Regist. XXXII (1846) 9; Grossh. Opr. (1949) 44 pp. "Fl. Cauc." IV (1950) 12 pp.
Syn. P. Wittmanniana auct. cauc. non Stev. pp. ; P. abchasica Mitz. Grossheim in "Fl. Cauc." II (1930) 92; Schipchin. in "Fl. USSR" VII (1937) 33; Kemularia Nathadse in "Fl. Georg." (1948) 6; P. tomentosa Kolak. "Fl. Abch." IV (1939)121.
Perennials. The stem is 80-100 cm high, with a branchy rhizome and not large root thickenings; at both sides leaves are nearly monotonously dark-green almost smooth, shining, bare or with sparse fine hairs at the lower side; leaf-lobes are oval, obovate, ovate with a long-narrowed base, sharpened or slightly blunt at the apex. The blossoms are broad patent, with pale-yellow, nearly whitish petals which are oblong-oval or oval, nearly not narrowed at their base, with a broad and short claw; anthers are yellow, filaments are yellow or flesh-coloured up to the middle, ovary is ovate with dense yellowish-tomentose pubescence, stigmas are nearly symmetrical-laminate, almost sessile or upon a very short style. The fruits are oblong-ovate, divirgent, yellowish-tomentose downy. The seeds are blackish-bluish, IV-VI.
Habitat. From the intermediary to upper montane belt, in the forest, upon margins.
Original area. Georgia, Abchazia.
Type. In London. It was described from samples grown in a garden of the Agronomic Society in Chisvik (the outskirts of London). The rhizomes gathered by Wittman in Abchazia were used in this case as a planting material.
Studied samples.
Georgia, Abchazia. The outskirts of Gagri; mnt. Mamdzishkha, 3, VII, 1912, I. G. Gusev!
Gorge of the river Bzibi, 13 km along the road to Ritse, slopes and bays of the inversional depression; 22. V, 1948, N. Purtskhanidze!
near Sukhumi; mnt. Apiancha. a margin of the beech-forest, 22, IV, V, 1902, Voronov! 20, V, 1953, T. Dumbadze!
Geographic type. Montane Kolchidsky. General distribution. Endemic.
Remarks. A history of this species is quite complicated. A yellow-flowered Caucasian peony had been firstly mentioned in 1842 by Fisher (the St. Peterburg Botanical Garden) in his letter to the famous English botanist Hooker, in London, who was informed that Hartwiss,( the director of the Crimean Botanical garden) had received from the count Vorontsov, from Abchazia, many interesting plants collected by Wittmann who travelled along the Caucasian Black sea coast, (afterwards, he became a garner in Odessa). Among the plants received, there was a yellow-flowered peony (along with Epimedium pinnatum Fisch.) which was propagated by Hartwiss throughout European States afterwards.
In 1846, in "Botanical Register", being issued by Lindley in London, materials about a new yellow blossomed peony named P. Wittmanniana Hartwiss, were published. This peony was described from samples grown in a garden of the Agronomic Society (London suburb.), and these samples had been obtained from the collection of Wittmann from Abchazia. In the diagnoses, it was indicated that the new peonies fruit was downy. Nevertheless, from a picture published the fruit looked naked.
It has happened so, that independently from each other, two authors described a yellow-blossomed peony found at different places and periods and had given to it the same name. A circumstance of this kind could not but mislead botanists, and some disagreement ensued among them subsequently in future. The majority of botanists acknowledged the authority of Steven and used, as a species name P. Wittmanniana Stev. Others botanists, especially from abroad, recognized the priority of Hartwiss, supposing the name Wittmanniana Stev. to be a synonym of the former or, at the most, considered it a variety with the name (var.) nudicarpa.
The author of the newest monograph about the genus Paeonia L. Stern (1946) has made a further step joining all yellow-blossomed peonies from Caucasus, excepting P. Mlokosewitschii Lomak from Lagodekhy, with P. Wittmanniana Hartwiss, to which P. Wittmanniana Stev. and P. macrophylla Lomak have been subordinated as its two i varieties by him.
We firmly object to the enlargening of species carried out in this way, especially, in case when such ancient representatives of the genus Paeonia L. as the Caucasian peonies are concerned. All Caucasian peonies with yellow flowers and growing at their natural habitats distinctly differ from each other with the complexes of morphological characters and the areas of distribution.
We quite agree with Stern (1946), who asserts that the priority concerning the species name and authorship belongs to Hartwiss, and that the peony with downy fruits and growing in Abchazia, but not in Talish or the North Iran, should be identified as P. Wittmanniana Hartwiss, and that the peony of Steven should be renamed, but not as a I variety, but as a self-dependent species.
It is quite obvious for us that Hartwiss (1846) and Steven (1848) described two different peonies from different places, Abchazia and Meskhetia, respectively, having given these peonies the same name.
We suppose that the identification of P. tomentosa (Lomak.) N. Busch from Talish with P. Wittmanniana Hartwiss is a result of an obvious mistake which is that in a box with plants sent to Hartwiss from Abchazia, there was a yellow-blossommed peony placed by a sample of P. Epimedium pinnatum Fisch, and the latter was known to grow in Talish only and, afterwards, botanists decided that the peony of Hartwiss should originate from Talish too; moreover, it was known that a yellow-blossommed downy-fruited peony had been found in Talish in 1888, and in the North Iran, in 1860.
But, at present, we are well aware of that, firstly, in the West Transcaucasus (Abchazia, Adjaria, etc.), the new peony species [Epimedium L. - Epimedium colchicum (Boiss) Trautv.] has been found and depicted, which is closely related, being almost indistinguishable, with Epimedium pinnatum Fisch. growing in Talish? secondly, in Abchazia, there is a yellow-flowered downy-fruited peony growing in Abchazia which is now under study by the prof. Mischenko and which is named P. abchasica Miscz. thirdly, it is well-known that Wittmann gathered plants near Tiflis, in Akhaltsikh, along the Black-Sea coast, and had never been in Talish. Hence, we must arrive at the conclusion that Hartwiss made his description of P. Wittmanniana in Abchazia and as to P. tomentosa(Lomak.) N.Busch. it grows in Talish and the North Iran; these two species noticeably differ from each other with the complexes of morpTioTogical features and with separated areas of distribution.
4. Paeonia tomentosa (Lomak.) N. Busch
Busch N. A. "Opred." Ranales Cauc. (1919); 7; Grossh. "Fl. Cauc." II (1930) 91; Schipczinsky, in "Fl. USSR" VII (1937) 26.
Syn. P. Wittmanniana var. tomentosa Lomak. in "Trudi Tifl. BG" II (1897) 283; Lipsky, "Fl. Cauc." (1899) 213; Schipczinsky in Not. Syst. Hort. Bot. Petrog." 2 (1921) 2 (1921) 44; P. Wittmanniana Busche, Aufzahl. Transcauc. Pers. Pfl. (1860) 8. Boiss., Fl. Or. (1867) 97 pp. ; Huth. in Engler Bot. Jahrb. 14. (1891) 266 pp. ; Stern, Monogr. in Jour. Roy. Hort. Soc. (1946) 56. P. Mlokosewitschii Grossh. (non Lomak.) "Opred." (1949) 44; Fl. Cauc. IV (1950) 13 pp. ; Rzazade in "Fl. Azerbadjana" IV (1953) 29. The picture in "Fl. Azerb." IV (1953) I sub P. Mlokosewitschii.
Perennials, having branchy rhizomes with radical thickenings; stem is long, 80-150 cm high, thick, firm, leaves are large, 6-8 cm long 3-6 cm wide, rough, dark-green, shrivelled, with distinguishably protruding veins at the lower side, grayish in result of pubescence formed by long hairs; leaf-lobes (leaflets) are ovate, obovate, elliptical with a shortly acuminate apex and a rounded nearly heart-shaped emarginate base. The blossoms are light-yellow, broad patent; sepals are downy from inside, petals are broad obovate, distinctly tapered at the base; anthers are yellow, filaments are violet or yellow, violet or reddish at the base; ovary is tomentose-downy, tangled with yellowish-grayish hairs; stigmas are flesh-coloured; filaments are purple as a whole or at the base only, in which case the upper part is yellow. The fruits are oblong ovate, 2.5-3 cm long, curvedly spreading, slightly covered with grayish-yellow hairs. The seeds are large, 7-8 mm long, covered by a net of large wrinkles.
Distribution. The upper montane belt, in forest, upon steep slopes.
Original habitat. Talish, a forest on mt. Nudus-Galasi.
Type. A forest on mnt. Nudus-galasi, 17,VI. 1894 L.Lomakin. The description was obtained from live specimens grown in the Caucasian department of the Tiflis Botanical Garden.
Specimens studied. Azerbaidjan: Talish. Forest near Nudus- Galasi 17,VI. 1894.L. Lomakin!
between Lyulyakeran and Orant 4. UII. 897 Alexeyenko!
Zuvant near the vill. Lerik, in forest 21. V. 1916.20.VII. 1917. Grossheim! 21.VIII.38. Rzazada!
west from Lerik. in shrubbery 4.V 1946. Ilyinskaya and Kirpichnikov!
Geographical type: Montane girkansky. General distribution. The East Transcaucsus. the North Iran.
Remarks. As far as a question of the Talish peony, P.tomentosa ( Lomak.) N. Busch is concerned we adhere to the point of view expressed by the authors of " Fl. USSR" ( 1937) and can't agree neither with A.A. Grossheim ("Opr." 1949. "Fl. Cauc." 1950) and P. Y. Rzazade ("Fl. Azerbaydjan" 1953), who identify this peony with the Lagodekhsky peony P. Mlokosewitschii Lomak. nor with Stern (1946) who considers it a synonym of P. Wittmanniana Hartwiss.
This species differs from the peony Mlokosewitschii quite distinctly with the heart-shaped or ovate lobes of rough leaves which are dark-green at the upper side and which ribs (vlins) protrude sharply, especially, at the lower side, and, also, with the stigma's and filaments colour, the petal's form and with the character of pubescence of ovary and fruits. A clear outlook on P. tomentosa (Lomak.) N. Busch. is illustrated by the picture printed in "Flora Azerbaydjana" (1953); unfortunately, a name of this plant was called incorrectly in this edition. As to the picture, published in the posthumous edition of "Fl. Cauc." (1950) by Grossheim, it should be regarded as fallacious, not typical for the Talishsky species, not to say about it having nothing in common with P. Mlokosewitschii (Lomak.).
The form of leaves and, especially, leaf-lobes make P. tomentosa (Lomak.) N. Busch. look somewhat alike with P. Steveniana Kem.-Nath. (P. Wittmanniana Stev.) that induced Lomakin to describe this plant as a variety and N. A. Busch to consider it a subspecies of P. Wittmanniana Stev. Hower this peony differs from the latter quite noticeably with the broad patent blossoms and, mostly, with the tomentose downy ovaries and fruits, and, also. with the separated areas of^rogafiation, that made N. A. Busch (1919) consider this plant a self-dependent species, afterwards. It is quite inadmissible not to distinguish these two species P. tomentosa (Lomak.) N. Busch from Talish and P. Wittmanniana Hartwiss (P. abchasica Miscz) growing in forest of Abchazia. Both of them have the only one common feature which is the ovary's and 11 fruits pubescence. Having green leaves at both sides, the latter species may be easily distinguished from P. tomentosa (Lomak.) N. Busch. which leaves are dark-green at the upper side and grayish at the lower side; moreover, the abchazian peony's leaf-lobes are tapered at the base; its pale yellow, nearly white petals are oval, almost rounded at the base; as to the Talishinsky species, its lateral leaf lobes are rounded at the base or nearly heart-shaped, petals are yellow, obovate, cuneately narrowed at the base. Also, these plants differ from each other with the separated areas of distribution.
So, it may be concluded that these two peonies are regarded as two independent species.
5. Paeonia Mlokosewitschii Lomak.
Lomakin in "Trudi Tifl. BG" II (1897) 282; Busch in Flora cauc. crit. III. 3. (1901) 14; Komarov, "Fl. USSR" VII (1937) 26; Grossheim, "Fl. Cauc." II (1930). IV (1950) 13 pp. "Opred." (1949) 44 pp; Stern in Journ. Roy. Hort. Soc. 68 (1943) 125. in Roy. Hort. Soc. (1946) 56; Kemularia-Nathadse in "Fl. Georgia; IV (1948) 5.
Perennials. The rhizome is woody, branchy with slightly thickened roots; stems are 80-100 cm high, branchy, densely-leaved; leaves are double-ternate, glaucescent at both sides, with very short, nearly unseen hairs at the lower side or bare at both sides; leaf-lobes are obovate, tapered at the base, rounded and shortly-bluntly-acuminate; veins are not coarse, thin,, slightly protruding. The blossoms are large, up to 10-12 cm wide, broad-patent; sepals are unequal, mostly concave and rounded, scarious-marginate at the brim, and, in less frequent cases, one of them is foliiform and much longer than others; petals are yellow, being dried out they get greenish at the brim often, ovate or obovate, slightly concave and slightly narrowed at base; stamens are with yellow anthers and filaments; ovaries are tomentose downy with yeltow curly hairs, ovate; stigmas are pink, sessile, almost symmetrical. The fruits are tomentose-downy, curved, spreading. The seeds are bluish-black.
Habitat. From the intermediary to upper montane belt, in forest, upon rocky slopes. Original area. Kachetia, the Lagodekhsky gorge, Ninigora.
Type. The description was obtained from samples grown in the Caucasian dept. of the Tiflis Botanical Garden, from rhizomes and seeds collected by Y. Mlokosewitsch.
Studied samples. Georgia. Kachetia, the Lagodekhsky gorge, a middle hill, 1901, Fomin! 1902, König! 5, V, 1935 Kapeller, Kemuaria-Nathadse! Ninigora! 1953, Kudigora! 1953, Kazarova and Dolukhanov!
Other parts of the Caucasus: Dagestan, Upper reaches of Avarsky Koys;
The basin of the Djurmut-or, left bank, 1958, A. Dolukhanov and N. A. Kazarova!
Geographic type. Montane Caucasian. General distribution: Caucasus. Endemic.
Remarks. It is a splendid species, and no disagreement exists between botanists about its self-dependance. Even Stern (1946) had no resolution quite enough to impinge on its self-dependance, despite him being so enthusiastic about joining all yellow-flowered peonies from Caucasus into one species.
With its bluish-grey, soft, smooth leaves, the lagodekhsky peony quite impresses imagination, looking so decorative, even at the time it has no blossoms. Also, its peculiar features are the pink colour of stigma, yellowish-curly pubescence of a nearly pyramidal ovary, and yellowish hue of filaments, that definitely distinguishes this peony from all other yellow-flowered Caucasian peonies (In "Flora USSR" these peonies are included into different serieses, and the Lagodekhsky peony is even considered to be in some proximity to P. oreogeton S. Moore, described in Mukden and propagating in Japan, China and the Far East).
As the matter of fact, among Caucasian peonies the P. Mlokosewitschii stands out separately, and it would be proper to establish a special series for it. But joining it with P. tomentosa (Lomak.) N. Busch from Talish and the North Iran, as it has been carried . out by A. A. Grossheim ("Opred." 1949 and "Fl. Cauc." 1950) and Rzazade ("Fl. Azerb." 1953) is quite irrelevant. We can't help commenting the pictures, published at -1 the above-mentioned editions and designated as P. Mlokosewitschii Lomak. which have . nothing in common with the Lagodekhsky peony and are depicting the Talishsky peony P. tomentosa, (the pictures have been taken from samples found in Talish, but not in Lagodekhy). The true picture of the Lagodekhsky peony is given by Stern (1. c.) and comparing it with the pictures from "Flora Azerbaydjan", it becomes obvious that there is a noticeable difference between them.
Sect. Paeon DC.
De Candolle. Prodr. I. (1824) 56 pp.; N. Busch in "Fl. Cauc. crit." Ill, 3 (1901) 8 pp. ; Stern, A Study of the Genus Paeonia in the Royal Horticalt. Soc. I (1946) 2.
Syn. Sect. Eupaeonia (DC.) Baillon in Adansonia 4 (1863) 56 pp. Sect. Palearcticeae, herbaceae Huth in Engler Bot. Jahrb. 14 (1891) 265 pp. Subsect Folioltae Stern I. c. 2. Subgenus Paeon (DC.) Seringe, Fl. jard. 3 (1848) 193.
Blossoms red, purple, pink, pinkish-violet; ovaries and fruits are tomentose-downy. The fruits are long, leaf-lobes are smooth-edged, broad.
Type. Paeonia mascula Mill.
Distribution. Montane countries of the South Europe and East Asia.
Series. Masculae Stern. Stern I. c. (1946) 67 pp. Series Corallinae Kom. "Fl. USSR" VII (1957) 28 pp. Sect. Corallinae Salm.-Dyck. in Hort. Dyck (1834) 365. 366 pp.
Blossoms red, purple or pink, leaf-lobes are obovate, oval or ovate and rounded at the base, tapered or rounded or nearly heart-shaped. Type. Paeonia mascula.Mill.
6. Paeonia Ruprechtiana Kem.-Nath. h. 1. combin. nova
Syn. P. triternata, forma coriifolia Rupr. Fl. cauc. (1869) 46; var. coriifolia (Rupr.) N. Busch in "Fl. Cauc. crit. Ill, 3, (1901-1903) 12; P. Kavachensis var. coriifolia (Rupr.) Grossh. in "Fl. Cauc." IV (1950) 12; P. caucasica var. coriifolia Rupr. in "Fl. USSR" VII (1937) 29; Kemularia-Nathadse in "Fl. Georgia" IV (1948) 6; var. viridifolia Kem.-Nath. 1. c.
Perennials with branchy rhizomes; the stem is usually 100-150 cm high, ribbed, robust. The leaves are leather-like, green, nearly monocoloured at both sides, lustrious, bare or sparsely hairy from beneath; leaf-lobes are broad and short ovate, oval, nearly rhomboid, shortly and abruptly narrowed at both ends. The blossoms are large, broad-patent, with reddish-pink petals; anthers are yellow, filaments are flesh-coloured, as a whole, or at the base only; ovary is oblong ovate with gold-shining tomentose pubescence formed by unequal hairs, stigmas are flesh-coloured, unsymmetrical with a second (one-sided) broad, wavy brim, nearly equal to an ovary height's half. The fruit is tomentose-downy.
Habitat. From the intermediary to upper montane belt. in shady forest.
Original area. The Kharagaulsky region, near the village Partskhnaly. The gorge leading to Lomis-mta.
Type. Imeretia, the Kharagaulsky region, near the village Partskhnaly, in the forest Tkhmelari 18. IV. 1916. I. Buachidze. The sample is kept in the Institute of Botany of Georgian Academy, in Tbilisi.
Studied samples. Georgia, Imeretia, the Kharagaulsky region, near the vill. Partskhnaly, in the forest Tkhmelari 18. IV. 1916, I. Buachidze!
Kartly. Borjom, II, VI, 1895 Radde!
outskirts of Borjom not far from the Baniskhevsky gorge II. V. 1918!
Borjom, in a forest, near the fortress Petristsikhe II. V. 1918 and 20. V. 1918!
descent from the ridge Dashiris-seri towards the Likansky gorge 8.VI.1920, V. Kozlovsky!
Geographic type. Kolchidia, General distribution. The Transcaucasus. Endemic.
Planta perennials. Rhizoma ramosum. Caulis plerumque 150, 200 cm, altus cylindraceus costatus validus glaber valde folious. Floria coriacea virida utrinque concoloria nitida glabra vel subtus sparse pilosa foliolis late ovatis ellipticus vel subromboideis basin et apicem breviter et abrupte attenaatis. Flores non late aperti magni roseo purpurascentes petala obovata basi cuneata. Anthera flava filamentis carneis in parte superiore flavis. Stygma lata dilatata purpurea. Ovarium fructusque tomentosi pilis flavis obsitis V.
Affinitis. Ab. P. caucasica N. Schipoz. follis utrinque viridis nitidis foliolis forma et petalis rosei bene differt.
Habitatio. In regione silvatica montim. Ainetum.
Typus. Georgia, Imerethi, Distr. Charagouli. Prope pag. Parzchnali. In silva Tchemlari (Ainetum) 18.IV.1916, I. Buatschidze.
Specimina vida: Georgia, Kartii, Distr. Borzhorni prope loc. Petriziche; fauces Banischevi II. V. 1918, inter Likani et monten Daqschiris seriin silva, 20.VI.1920 B. Kozlowsky.
Area geogr. Caucasus, Georgia. Species endemica.
Remarks. This species attracts one's attention with its almost monocolored at both sides, shining leaves which get lustrous after being dried out, and with its nearly rhomboid or broad and short ovate leaf-lobes shortly tapered at both ends. And with all above-mentioned, this species differs quite distinctly from other known red-flowered peonies.
Yet in 1869, studying collections of Radde from Borjom, Ruprecht described a new form of peony named P. triternata, forma coriifolia Rupr; and in 1902, assuming the leather-like consistency of leaves and absence of wax-like bloom on them to be constant characters for the genus Paeonia L., N. A. Busch defined this plant as a variety.
All subsequent botanists adhered to the opinion of N. A. Busch, and only a name of the basic species changes, depending on comprehending of it by researchers.
After Radde, for long time, nobody gathered this plant. But in 1918 and 1920, it has been found by V. L. Kozlovsky nearby the outskirts of Borjom, near the fortress Petristsikhe, in the shady gorge, where this plant was collected by Radde obviously earlier, and in the Baniskevsky and Likansky gorges.
Earlier in 1916, I. Buachidze collected peonies in the "shady gorge" leading to Lomis-mta, at the border with the Borjomi region, in the Kharagaulsky region (in past, it was included in the former Shorapansky dist.). Afterwards, (in 1946), this peony has been described by us as a variety named P. caucasica var. viridifolia Kem.-Nath.
As it was found out later, these two varieties are identical. Besides, some other characters have been discovered which make these two peonies differ from the Caucasian red-flowered peony P. caucasica Schipcz., which are such as, the short broad ovate or nearly rhomboid, abruptly-shortly narrowed at both ends leaf-lobes; the obovate, cuneately tapered at the base, petals; the broad patent blossoms; the gold-shining tomentose pubescence upon ovary, etc.
At present, an area of distribution (habitat) of this plant has been found out and, taking into account a complex of the above-mentioned morphological characters, this peony may be recognized for a self-dependent species, for which I give a name in honour of the famous botanist, academician Ruprecht, who distinguished it from the well-known Caucasian red-flowered peony, implying by words "haec forma coriifolia forsan nova species est" a possibility for his form to be acknowledged for a self-dependent species in future.
7. Paeonia caucasica Schipcz.
Schipczinsky in "Fl. USSR" VII (1937) 28; Kemularia-Nathadse in "Fl. Georgia" IV (1948), 6.
Syn. P.corallina var. caucasica Schipcz. in Notul. Syst. Horti. Bot. Petrop. II (1921) 45. ,. P. corallina var. triternata (Pall.) Boiss. Fl. Or. (1867) 97 pp. ; P. corallina subsp. triternata (Pallas) N. Busch in Fl. cauc. crit. 3.III.(1901-1903) 10 pp. ; Ledeb. Fl. Ross. I (1842) 73 pp. ; Ruprecht. Fl. Cauc. (1869) 44 pp. ; Stern in Journ. Roy. Hort. Soc. 56 (1931) 74 pp. ; P. kavachensis Grossheim, Fl. Cauc. IV (1950) 12 non Aznavour; P. daurica Stern in Journ. Roy. Soc. (1946) 70 pp. . XXIV
Perennial. The rhizome is palmate-ramificating with slightly swollen root thickenings; stems are from 50 to 100 cm high, robust, branchy; leaves are double-ternate, dark-green from above, grayish, in result of pubescense, from beneath or glaucescent at both sides or only at the upper side; leaf-lobes (leaflets) are oblong-obovate or oval, shortly acuminate. The blossom is large, 3-5 cm in diameter, with red petals, yellow anthers and red or violet filaments; ovary and fruits are tomentose-downy with yellowish or grayish hairs; fruits are large, curvedly spreading. Seeds black and red; unripe seeds are red, fleshy.
Habitat. From the intermediary to upper montane belt, in forest, upon margins. Original habitat. The West Transcaucasus. The type has not been preserved.
Studied samples. Georgia, Abchazia. By mnt. Chipsira 3.X. 1893, Albov!
Near the vill. Yuryevskoe, Sukhum, IV. 1900. Voronov!
Near the vill. Tsebelda, 1956, Dumbadze!
Bridsky ridge 1948, Purtskhvanidze!
Svanetia. Lentekhy. 28. VII. Some and Levye!
Racha-Lechkhumi; a gorge of the river Rion, mnt. Konchkhu by the vill. Achariskhidi (Achara) 12.VII.1929, Kemularia-Nathadse!
Between the vill. Mikotsminda and Abanoyety, in forest 22.IV. 1935 M. Sokhadze!
Megrelia. Mut. Ochakhuye, a forest margin at the upper belt 22.VII.1954, Kemularia-Nathadse, Mandenova, Kutateladze!
between the vill. Leshamge and mnt. Barkha, in forest 15.VII.1954. M. Dolidze and I. Mikeladze!
Imeretia. Kvirili, an oak forest IV. 1916, Lomakin!
Sakolovsky ridge beyond the vill. Gavati, in forest 20.VII.1927!
in a forest between the vill. Gelati and Godogani 26.VII.1926, 22.IV.1930, Kemularia-Nathadse!
the Tkibulsky region, beyond the vill. Akhalsopeli, upon mnt. Tsintskala, 4. VI. 1948. A. Kutateladze!
Adjaria. Upon the ridge Khanii, 28.VII. Somye and Levye!
Kartly. The outskirts of Tbilisi, in forest, IV. 1844 Hohenacker!
Martkobi, abundantly in forest, 15.IV.1861, Ruprecht!
Borjom 1876, Medvedev, 28.VII.1888 Akinfiev! II.V.1918, Kodjori!
in forest 26.V.35, Barnabishvili! Zedelmeyer!
Saguramo 7.V.1920, Kafiev! 12. V. 1929 A. Kolakovsky! 23. V. 1920., Zedelmeyer!
Kakhetia, Kakabeti, in forest Dvrina 8. VI. 1914, Kanchaveli!
Kardanakh, shady places 4. V. 1900, Grinevetsky!
Azerbaydjan. Lenkoran, in forest. May of 1838, Hohenacker.
Other parts of the USSR: RSFR, the Stavropolsky region, Klukhorsky distr.
Geographic type. Montane Caucasian. General distribution. Caucasus. Endemic.
Remarks. In the Caucasian botanical literature our red-flowered peony was mistaken for P. corallina Retz. (the South-European species) or for P. triternata Pall. growing in the Crimea, for too long. Just recently. Grossheim designated this plant by the name P. kavachensis Aznav. applied to a peony from the Kavachsky region in Turkey (1949, 1950). N. Busch (1901) denied existence of the typical form P. corallina Retz. with peculiar narrow lanceolate leaf-lobes in Caucasus and identified our Caucasian peony with the Crimean peony, considering the latter a subspecies of the Europen species.
N. V. Schipczinsky (1921) separated the Caucasian peony from the Crimean peony, considering the former a variety of the European species P. corallina Retz., and in 1937 he declared it as a self-dependent species named P.-caucasica Schipcz.
In 1946, Stern proclaimed the Caucasian peony to be a synonym of the Crimean P. daurica Andr. Consequently P. corallina Retz. and P. kavachensis Aznaur. were assumed by him to be synonyms of P. mascula Mill. Stern separated them from our peony that was quite correct, but identified the Caucasian peony with the Crimean species, that was a mistake.
Undoubtedly, P. caucasica Schipcz. is closely enough related with the Crimean peony, but from which it differs quite noticeably with the yellowish pubescence of ovaries and fruits, with flesh-colored stigmas, large leaf-lobes which are glaucescent at the lower side only and with more vigorous, high stems. Recognized by the name P. caucasica, our Caucasian red-flowered peony is not homogeneous throughout the area of its propagating. Yet in 1869, Ruprecht from the Borjomi region marked out by the name P. corallina forma coriifolia Rupr. a peculiar form of peony with nearly leather-like lustrous green leaves. In 1948, we have presented 4 varieties, such as var. viridifolia, var. glauca, var. pubescens and var. gigas. The first of them occurred to be identical with the form of Ruprecht, which is defined by us as a self-dependent species (see above P. Ruprechtiana); the second and last varieties are to be joined with the typical form, for which the samples from the West Transcaucasus are recognized, as it was indicated in "Fl. USSR" (1937).
Besides, in result of our researching, we have reached a conclusion that the pink-flowered peony from the Lagodekhsky gorge is a self-dependent species which is submitted below with the name P. lagodechiana Kem.-Nath.
As to the red-blossommed peonies mentioned by old authors and growing in Armenia and Talish, we could not find out anything about them. In "Flora of Armenia" (1954) peonies are not indicated at all, and in "Flora of Azerbaidjan" (1953) only P. tenuifolia L. and P. Mlokosewitschii Lomak. are mentioned, and the latter is mistaken for P. tomentosa (Lomak.) N. Busch.
Some indications in "Armenia rossica" given by Schovits and his herbarium sample preserved in Leningrad pertain to some regions of Georgia, Trialetia or Kartly, probably.
As to the red-blossommed peony, indicated by Stern, as P. mascula Mill. and found in Azerbaydjan, not far from Lenkoran, it remains unclear what it may look like. ("Russia; Azerbaydjan, near Lenkoran, Hohenacker (K)").
8. Paeonia lagodechiana Kem.-Nath. h. I. species nova
Perennial, 50-100 cm high, plant with a branchy rhizome having spindle-like root thickenings; the stem is ribbed, densely leaved; leaves are double ternate, glaucescent at both sides or only from beneath, smooth with slightly protruding veins, bare; leaf-lobes are smooth-edged, oblong-elliptical, at both sides or at the apex they are shortly narrowed. The blossoms are pink, broad patent, with obovate slightly concave petals; anthers are orange, filaments are purple; ovary is yellow, tomentose downy; stigmas are flesh colored, almost symmetrical. The fruit is tomentose-downy.
Habitat. The intermediary montane belt, in forest.
Type. The lagodekhsky reservation. Ninigora, in forest 10. V. 1953 A. Kazarova and A. Dolukhanov.
Studied samples. Georgia, Kachetia. the Lagodekhsky gorge 5.V. 1935 Kemularia-Nathadse P. Tsagarely.
Planta perennis, plerungue 50-100 cm. alta. Rhizoma crassum ramosus, radix fusiformis. Caulis cylindraceus vix angulatus glaber ramosus dense foliosus. Folia biternato secta longe petiolata superne viride subtus vel utrinque glauca, foliolis ellipticis vel oblongo-elliptics apice acutis basin versus breviter attennatis glabris nervis non prominentibus. Flores late aperti rosel petala vix concava obovata basi cuneata. Stamina flava filamentis stigmaque carneis. Fructus tomentosus, IV-V.
Affinitas. Ab P. caucasica N. Schipcz. floribus roseis late apertis petalis foliolisque forma bene differt.
Habitatio. In regione silvatica montium et margines silvarum.
Typus. Georgia. Rauces Lagodechi. loc. Ninigora 10. V. 1953 A. Kazarova et A. Doluchanov.
Specimina vida: Georgia. Fauces Lagodechi in silva 10.V. 1935 P. Zagareli; 5. V. 1936 Kemularia-Natadze.
Area geogr. Caucasus. Species endemica.
Remarks. It is a very effective, ornamental plant. With its tender glaucescent leaves, it resembles P. Mlokosewitschii Lomak., nevertheless from which it noticeably differs with its pink larger broad-patent blossoms, stigma's, anthers and filaments colour and leaf-lobes form, also. As to the latter character, it is obovate, blunt at the apex and long-tapered at the base for P. Mlokosewitschii Lomak. From P. caucasica N. Schipcz. it differs with its pink broad-patent larger blossoms, with petals and leaf-lobes form. Till present it is known to have been found in the Lagodekhsky gorge only.
9. Paeonia triternata Pall.
Pallas in Nova Acta Acad. Petrop. X (1975) 312; DC. Prod. I (1824) 65; Ledeb. Fl. Ross. I (1842) 73; Rupr. Fl. Cauc. (1869) 44; Lipsky Fl. Cauc. (1899) 213; Somm. et Lev. Enum. (1900) 29; Komarov and Schipczin. in Fl. USSR. VII (1937) 28;
Syn. P. corallina var. triternata Boiss. I. (1867) 97; Akinf. Fl. Centre Cauc. (1894) 50; Schmalg. Fl. Sredn. Juzn. Ross. (1895) 32; P. corallina ssp. triternata N. Busch in Fl. cauc. crit. III. 3,(1901) 10, P. corallina var. Pallasi Huth. Monogr. in Englers Bot. Jarhb. 14 (1891) 267; P. corallina Retz. in M. B. Fl. taur-cauc. 2 (1808) 10; P. corallina Grossheim Fl. Cauc. II (1930) 91; P. daurica Andrews. Bot. rep. 7 (1807) 486; Anders. Mag. (1812) 1841. Stem, Monogr. Royal Horticult. Soc. (1946) 70; P. taurica Grossh. Opred. (1949) 44, Fl. Cauc. IV (1950) 12.
Perennials. The rhizomes are thickish, robust, with oblong-lengthened root thickenings, which are sessile usually or upon short pedicels, stems are 50-100 cm high, often tortuous, thin ; leaves are thrice ternate, naked, bluish-gray at both sides; leaf-lobes are small, rounded-ovate or nearly heart-shaped at their base, nearly leather-like; blossoms are broad-patent, with purple, oval or obovate petals; filaments are yellow; ovary is with pink tomentose pubescence; stigmas are pink. The fruits are with pinkish-brown pubescence, spreading; seeds are dark-blue, IV-V.
Habitat. In the montane oak and beech forests, upon open slopes.
Original area. The Crimea.
Type. The description has been made in the Crimea. Preserved in Leningrad.
Studied samples. Prichernom., Anapa, 13.VI.91!
Novorossiysk, 22.IV.1889 with flowers, 6. V. 1892 Lipsky!
50 miles south from Novorosiysk IV-1897 Akinfiev!
Krasnaya Polyana (the middle stream of the rivers Mzimti, Achipsou) Albov!
Geographic type. Crimean montane. General distribution. The Crimea, Caucasus.
Remarks. This plant has been firstly reported by the academician Pallasi in 1795, from the Crimea and named P. triternata Pall. But the latm diagnosis has been given by De Candolle in 1824, and for this reason some botanists prefer to use the name P. daurica Andrews for peony described in 1807 and found in the Crimea too (Stern 1946, Grossheim 1949, 1950).
We suppose that it would be correct to retain the name for this Crimean peony given earlier by Pallasi, and according to the opinion of De Candolle, Komarov and Schipczinsky, designate it P. triternata Pall. Not to say about that the name given later may mislead future botanists.
Considering a question of this species being propagated beyond the borders of the Crimea, and, consequently, of its presence in Caucasus, we have been looking through quite many herbarium samples of red-flowered Caucasian peonies and never met any typical representative of the P. triternata species, having such peculiar features as the pink or pinkish pubescence of ovaries and fruits, pink stigmas and small rounded or heart-shaped at its base leaf-lobes at both sides of gray-blue leaves with wax-like bloom upon them. Hence, it may be concluded, that the P. triternata Pall. does not grow in Caucasus; nevertheless, we present it here, as a representative of "Flora of the USSR“ for the Novorosiysk region, and submit here more accurate data on its distribution area, using partly materials of Busch (L.c.) and partly the herbariums samples, having been studied by us in the Botany Institutes of Leningrad and Tbilisi and which were quite scarcely represented, it should be mentioned also.
Sect. Sternia Kem.-Nath. comb. nova. Syn. Subsect. Dissectifolia Stern Study of the Genus Paeonia in the Royal Horticult. Societ. I (1948) 8 pp.
Folia bi vel triternata , foliolis profunde lobatis, dentatis, triternato v. pinato dissectis. Corollia purpurea v. purpureo-rosea. Ovarium fructusque brevi, tomentosirarissime glabri.
The leaves are biternate with lobes being deeply incised or pinnately dissected into lanceolate or linear lacinules ; corolla purple, dark-red, blackish purple or reddish-pink, ovaries and fruits are tomentose-downy, short, ovate or oval. Type P. officinalis L. emend. Willd.
Series Hybridae Kem.-Nath. Series nova. Syn. Series Dentate Komar. in Fl. USSR VII (1937) 33 pp. Anomala group Stern I. c. pp.
The leaf-lobes are triternate pinnately dissected into narrow lanceolate lacinules. Foliolia triterato pinnatisecta, seqmentis (lobis) anguste lanceolatis. Type P. hybrida Pall.
10. Paeonia Majko N. Ketzch.
N. Ketzchoweli in Notul. syst. ac geograph. Inst. Bot. Tiflis. fasc. 21 (1959) 17.
Perennials. The rhizome is branchy, nearly horizontal, with fusiform root thickenings; stem is high, densely leaved, with lasting lower leaves; leaves are green, thrice pinnately incised or dissected into narrow lanceolate 5-10 mm wide lacinules; leaf lobes of leaves at the root base are broader than for the upper leaves; flowers are large, pinkish-reddish-violet, broad patent; ovary and fruits are with dense tomentose pubescence formed by red hairs; fruits are short, oval or ovate.
Habitat. Foot-hills, margins of the oak-hornbeam forests.
Type. Georgia. Kartly. Between the vill. Lamitskali and Igoti, 16.V.1958, N. Ketskhoveli.
Studied samples. Kartly. The vill. Lamistskali, V.1949, Lumbadze.
Remarks. It is a very decorative, beautiful plant. With its leaves and pink flowers, it resembles P. hybrida Pall., growing on steppe meadows, open grassy or rocky slopes of hills in the West Siberia (Irtish, Altay) and the Middle Asia (Pamir, Altay, Tyan-Shyan), and having been described in Barnaul. Moreover, it has some resemblance with the peony, named as a hybrid. P. tenuifolia P. triternata which has been described by V. P. Maleev in 1937, in the Crimea ("Sovetskaya Botanica" 1937). Maleev asserts that his peony is a hybrid by reason of it having been found by V. F. Vasilyev by around the vill. Uzundja, upon the ridge Dara-Bayr, where these two Crimean peonies P. tenuifolia and P. triternata grow together.
Until the question of hybridity of this peony is solved through experiments, we accept the Maleevsky peony as a geographical race, just as P. Majko Ketzcch. and P. hybrida Pall, naming it P. Maleev Kem.-Nath. in honour of Maleev, and join all these peonies in the series Hybride Kem.-Nath.
It is necessary to point out at these peonies remote resemblance with P. anomala L. described in Siberia and typical for "the forest zone, exclusively" of the Europe part of the USSR. West and East Siberia and Middle Asia. The latter species is included into the series Dentatae Kom. ("Fl. USSR") along with P. hybrida Pall.; so Stern was quite right having established a series for it with the name Anomala group, which includes P. Veitchii Lynch, P. Beresowskii Kom. and P. Woodwardii Stapf., all of them from China.
Obviously, all above-mentioned species had a remote mutual ancestor which had been growing in forests of the tertiary period and being more close to P. anomala. As environmental conditions were changing and forest areas decreased, steppes began forming and new species of peonies started arising, from which the youngest species should be considered P. tenuifolia.
Series Fissa Kom. Fl. USSR. VII (1937) 31; officinalis group Stern I. c. pp.
Leaves double-thrice ternate or pinnate, multifid with narrow lobes. Folia bi-vel-tri-ternata vel bipinnata lobis anquste linearis.
Type. P. tenuifolia L.
11. Paeonia tenuifolia L.
Linneus Sp. pl. (1763) 748; Ledeb. Fl. Ros. I (1842) 73; Boiss. Fl,. or I (1867) 98; Schmalhausen, Fl. Sr. i Juzn. Rossii, I bvfr(1895) 73; Lipsky, Fl. Cauc. (1899) 213; N. Busch in Fl. Cauc.crit. 3.III.(1901) I; Komarov and Schipczinsky in Fl. USSR, VII (1937) 34. Stern, in Roy. Hort. Soc. (1946). Grossheim, Opred. (1949) 44; Fl. Cauc. IV. (1950) 12; Rzazade in Fl. Azerbaidj. IV (1953) 30.
Perennials. The rhizomes are with oblong, cone-shaped root thickenings upon short pedicels. The stem is simple, 10-15 cm high, densely leaved; leaves are with very narrow nearly filiform 1-2 mm wide pendent lobes. The blossoms are large, 3-5 cm in diameter; petals are dark-red, obovate, rounded at the apex and base; anthers are yellow; filaments are reddish; ovary and juvenile; fruits are with dense red tomentose pubescence. The fruit is short-ovate or oval, slightly spreading aside, with brownish-red pubescence; seeds are brown-black, shining, IV, V.
Habitat. The steppe zone, among shrubbery.
Type. Preserved in London. Original area. Ukraine, the outskirts of Taganrog.
Studied samples. Armavir region (North Caucasus) 8.V. 12.V. 1889, 12.IV.I 1891.!
Nevinomiskaya 15.V. 1889 Lipsky!
Sergievskoe VI. 1893!
Kislovodsk 2.VI.1893!
near Pyatigorsk 15.IV.1897 Akinfiev!
the river Kuban, Radde!
Kuban, Veliaminovskoe 25.IV.1907 N. Busch and V. Klopotov!
Prichernomorye. Near Anapa, IV. 1900, 0. Dekleiz and Y. Voronov!
Novorossiysk, montane meadows above the cement factory, 23.IV.1902 A. Fomin!
Georgia. Kartly. The Gorysky region, the outskirts of the vill. Samtavisi 16.IV.1918 Volochanetsky!
Mikhetsky region near the vill. Tsilkani 6.IV.1958 and 6. VI. 1949 Dumbadze!
Kizik., Schiraki, the Gorge LekisO-Khevi 20.IV.1958 Kakheladze and Gavrilenko!
Azerbaidshan. Karabulakh V. 1902. Schneider from the herbarium of Medvedev!
Geographic type. Pontean. General distribution. The middle Europe, the European part of the USSR, the Middle Dnjepr; Volga-Don, the Crimea, the down stream of Volga.
Remarks. Describing P. tenuifolia L. found in the outskirts of Taganrog, Ukraine, Linneus characterized it as having leaves manifold dissected into narrow, linear lobes. Willdenow added a new character which is the purple hue of ovarie's pubescence and which has been noticed and pointed out at by Pallas just too long ago, in 1773. Afterwards, these characters were ignored by the most of botanists, excepting the authors of "Fl. USSR" only.
Nevertheless, the hue of ovarie's pubescence is a constant quite important taxonomic feature for the genus Paeonia L., being a well distinguishing character for the related species, such as P. tenuifolia L., Biebersteiniana Rupr. , tenuifolia L. and P. carthalinica Ketz.
All these species are very closely related with each other composing one evolutinal series lFissae Kom., which should be named after one of its species, that might have been P. tenuifolia L., but taking into consideration priority demands, we retain the name given by the academician Komarov previously.
As to the distribution of P. tenuifolia in Caucasus, opinions of botanists diverge considerably. Most of them suppose that this species is typical for steppe regions of the North Caucasus, Transcaucasus, including Georgia, Azerbaydjan and Armenia (Grossheim 1930, 1950). In "Fl. USSR" the P. tenuifolia is indicated as growing in Predcavkaskye only. Although the question about what a peony with leaves dissected into narrow lobes and named P. tenuifolia L. from TransCaucasus, according to some old authors (e.g. Boisser, 1867, et al), does just happen to look like, is silenced somehow.
In the first edition of "Fl. Georgia" (1948) we expounded an opinion that in Georgia and Transcaucasus, as a whole, the P. carthalinica ketzch grew instead of the P. tenuifolia L., the both being closely related with each other; nevertheless, new data obtained from collections by researchers from the Institute of Botany (N. Kakheladze, B. Gavrilenko, Kakulia, M. Sakhokia, M. Sokhadze, Kebadze) and from the Botanical Garden (T. Dumbadze, B. Serdyukova), and our own observations made us reject our previous opinion and at present these two species are submitted here in our present work.
As to the presence of P. tenuifolia L. in other republics of Transcaucasus, on a base of data available in "Flora Azerbaydjana" (volume IV, 1953) it may be concluded that this plant may grow in Dzegam, Pirchevan, Nakhichevan. Also, the indications by old authors in concern to P. tenuifolia L. growing in Armenia and which were based on the herbarium materials of Szovitz (herbarium samples with labels "Armenia rossica") and A. A. Grossheim's data "Fl. Cauc." 1950, v. IV), have not been corroborated by recently obtained data by A. L. Takhtadjan. So, in "Fl. Armenia" (v. I. 1954) the genus Paeonia L. is not mentioned at all.
12. Paeonia carthalinica N. Ketzch.
Ketskoveli. The basic plants types of Georgia (1935) 20 (descr. georg.) in Notul. Syst. ac georg. Inst. Bot. Tifl., 21 (1959) 18; Kemularia-Nathadse in Fl. Georgia IV (1948) 7. Syn. P. tenuifolia auct. cauc. pp.
Perennial. The rhizomes are branchy with round, nearly global or round-ovate cone-shaped root thickenings; stems are simple or branchy, 30-100 cm high; leaves are manifold dissected into long narrow-linear unpendent lacinules with a width 5-10 mm; upper cauline leaves are wrapping a blossom. The blossoms are 5-7 cm in diameter, with dark-purple petals which are fusiformly tapering at the base; anthers are yellow; filaments are dark-purple; ovary is ovate with dense yellowish-tomentose pubescence. The fruits are ovate, almost upright, with yellowish-tomentose and less frequently red pubescence; getting ripe, the fruits become yellowish-grayish or gray downy. The seeds are oblong, blackish-brownish, III.IV.V.
Habitat. Foot-hills, in light oak-hornbeam forests or upon margins, mesophyllous variants of the prickly-shrubbery formation.
Original area. Georgia, Kartly.
Type. Kartly, between the villages Igoiti and Lamiskana 15. V. 1935 N. Ketskoveli.
Studied samples. Kartly, the Goriysky region. Mukhrani beyond the vill. Dampalo 15. V. 1935 Ketskhoveli, Kemularia-Nathadse!
Metskhetsky region, beyond the vill. Tsilkani 12. V. 1949 Dumbadze!
vill. Igoiti 15. V. 1958, 4. V. 1959 N. Ketskhoveli!
Shiraki. Between the vill. Keda and mm. Shavimta 19.VII.1927 Z. Kanchaveli!
the Peony gorge 4. V. 1955 Kakulia, Sokhadze, Kebadze.!
Remarks. It is very closely related with P. tenuifolia L. from which it differs with broader leaf-lobes and grayish pubescence of ovaries and fruits. With the above-indicated features, the Kartlinsky peony resembles P. Biebersteiniana Rupr. growing upon Stavropol hills; the latter species differs from the Kartlinsky species with grayish leaves which lobes are covered with hairs growing in rows along veins at the upper side.
Possibly, the Kartlinsky and Stavropolsky peonies are identical, representing one species with a disrupted area of distribution. Nevertheless, to solve this question new collections and, mainly, observations in natural conditions of growing, are needed.
Till then we prefer to consider these plants self-dependent species joined in one evolutional series.
13. Paeonia Biebersteiniana Rupr.
Ruprecht. Flora Cauc. (1869) 47; Trantvetter Incr. (1882) 61; Lipsky, Fl. Cauc. (1899) 214; Komarov and Schipczinsky in Fl. USSR VII (1936) 35; Grossheim, Opred. (1949) 44. Fl. Cauc. IV (1950) 13.
Syn. P. tenuifolia var. Biebersteiniana (Rupr.) N. Busch in Fl. cauc. critic. Ill, 3 (1901-1903) 9; Grossheim Fl. Cauc. II (1930) 91; P. tenuifolia v. hybrida (Pall.) Lipsky Fl. (1898) 235; P. tenuifolia Rupr. Fl. cauc. pp. ; MB. Fl. taur.-cauc. II (1808) 10 pp.
Perennial, its rhizome has oblong cone-shaped root thickenings; stems are 20-70 (100) cm high, simple or less rarely branchy at the upper part; leaves are thrice ternate-incised into narrow-linear, 3-10 mm wide, short, unpendent lacinules, which are naked or covered with short hairs, growing in rows along veins at the upper side. The blossoms are middle-sized, with bright-red petals, yellow anthers, red, less frequently yellow filaments; ovary is ovate, tomentose downy; stigmas are red; fruits are upright or slightly spreading. Seeds .....V.
Habitat. Over steppe slopes, among shrubberies. Original area. The outskirts of Stavropol.
Type. Outside of Stavropol, 1867, Becker, Kept in Leningrad.
Studied samples. North Caucasus, Stavropol 1827, Steven! Marshal-Bieberstein! 1867 Becker! 28, Lipsky!
Georgievsk, Wilhelms!
Chechenia, Grozny 5. V. 1890 Lipsky!
Stavropolskij Kraj, Near t. Pyatigorsk V. Hochenacker!
Near Zakan-Yurtovskaya among shrubberies, Kuznetsov!
Geographic type. Montane steppe. General distribution. The North Caucasus.
Remarks. In respect to the question of self-dependance of this species, the opinions of botanists diverge quite significantly. Most of them consider it a variety or form of P. tenuifolia L. N. Busch considers this plant a transitional form from P. tenuifolia L. to P. anomala L. growing in Siberia Denying independence of this species, he writes: "The var. Biebersteiniana (Rupr.) N. Busch can be regarded as a self-dependent species by no means, because it differs from P. tenuifolia L. with broader leaf-lobes only".
Huth (1891) took P. Biebersteiniana for P. tenuifolia L. and pointed out at P. anomala L. growing in Siberia, as a species typical for Stavropol. A fallacy of this kind was commented by N. Busch (1901).
Stern (1946) considers this plant a "tantum nomen", only, and places it in the separate list "Species non satis notae", as not being adequately studied.
The self-dependence of this species is indicated by no less authorize botanists, such as Ruprecht, Trautvetter, Lipsky, Komarov (Fl. USSR, 1937).
As it seems to us, a disagreement of this kind is a result of different concepts about species volume being used in every case. But the main reason is the absence of well-furnished herbariums and proper interest for these species. Occasional desultory gatherings of the plants were carried out, without taking into account ecologic and geographical characteristics in this case. Only on a base of massive collections and regular observations in natural conditions, it is possible to reveal differences between so closely related species, as P. tenuifolia L., P. Biebersteiniana Rupr. and P. carthalinica Ketzch.
As indicatory features for comparing these species, such the characters are mentioned as follows: the larger width of leaf-lobes of P. Biebersteiniana Rupr. in comparison with P. tenuifolia L. and the row of hairs at the upper side of grayish leaves. But if the ovary's and fruit's pubescence made up by red hairs is taken into account for P. tenuifolia, as it was pointed out by Gmelin, Willdenow in "Flora USSR" previously, than a complex of characters needed for recognizing the species will be obtained in this case.
Besides, these plants have different areas of distribution. Just as the outskirts of Stavropol including hills (the Stavropol plateau) are known to be the only area for P. Biebersteiniana Rupr. distribution, so this peony may be considered an endemic species for this region (see "Fl. USSR"). So, having been described in the steppe zone of Ukraine (the outskirts of Taganrog), P. tenuifolia L. is known to be propagating throughout steppe regions of the South European part of the USSR, North Caucasus. Transcaucasus and the Middle Europe.