The Caucasian Representatives of the Genus Paeonia L.

L.M. Kemularia-Nathadse, Trudy Tiflis. Botan. Sada 1961

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Chapter V

Considerations pertaining to the Caucasian species of the genus Paeonia L.

As far as comprehending of the Caucasian representatives of the genus Paeonia L. is concerned, we have some contradictions with Stern, the author of the newest monograph about peonies (1946). As it seems to us, he has not made quite clear for himself the problem concerning the Caucasian peonies, that is quite explicable, since he had no materials about these plants, particularly, about yellow-blossommed peonies. Stern writes that herbarium samples of yellow-blossommed peonies are very rare; most of them come from the Tiflis Botanical Garden, and the only herbarium sample of wild yellow-blossommed peonies seen by him in the Britain Museum was collected in mountains of Elbrus, in the North Iran, to the west from Astrabad, and, besides, this sample happened to be not complete.

So, Stern did not acquaint himself with any Caucasian peony's herbarium samples from their natural habitat, and all his considerations were based on literature data provided by Russian botanists in different periods, according to his own account. At our disposal we had live specimens growing in the Tiflis Botanical garden and the department of live plants of the Botany Institute of Georgian Academy Science, not to say about herbarium samples of our own and other persons collections; -most of these plants have being studied by us in environmental conditions, also.

From Caucasian yellow-flowerd peonies, Stern recognized, as self-dependent species, P. Mlokosewitschii Lomak. and P. Wittmanniana Hartwiss ex Lindl. with two varieties nudicarpa (P. Wittmanniana Stev.) and macrophylla (P. macrophylla (N. Alb.) Lomak.). As to P. abchasica Miscz. and P. tomentosa (Lomak.) N. Busch. he considers them synonyms of P. Wittmanniana Hartwiss ex Lindl.

Stern is right acknowledging the priority of P. Wittmanniana Hartwiss and considering P. abchasica Miscz. a synonym of the former peony. In fact, P. Wittmanniana Hartwiss has been described earlier, and P. Wittmanniana Stev., being described later, should be renamed, but not as a variety, but as a self-dependent species.

Also, we consider P. macrophiylla (N. alb.) Lomak. a self-dependent species (details are given in references about species). According to N. Busch, naked fruits occur to be a good character for identifying species and, as a result, our Caucasian peonies with naked fruits (and ovary's form, too) evince more affinity with the Far-East yellow-flowered peonies, than with our downy-fruited peonies; as it has been found out, all peonies with yellow and yellowish blossoms from the East Asia and Japan (P. Lutea, P. Delavayi, P. lactiflora Pall., P. japonica Myabe et Takeda, P. oreogeton S. Moore, P. trollioides Stapf., P. obovata Maxim., P. Willmottiae Stapf.) have naked ovaries and fruits, so that this character occurs to be more primitive and, as a result, ancestral.

The P. tomentosa (Lomak.) N. Busch has been described by A. A. Lomakin in Talish, as a variety of P. Wittmanniana Stev., that is as a peony with naked fruits from Akhaltsikh; Lomakin and other subsequent botanists knew nothing about a downy-fruited peony from Abchazia. Just at present we know quite well that two different peonies from different places with names P. Wittmanniana Hartwiss and P. Wittmanniana Stev. have been described by two authors on a base of collections of the same collector (see references about species) and, as a result, the yellow-flowered downy-fruited peony from Talish could not be a variety of the bare-fruited peony from Akhaltsikh, and therefore N. A. Busch was quite right separating the Talishsky peony from P. Wittmanniana considering the former a self-dependent species. It is incorrect to identify it with P. Wittmanniana Hartwiss from Abchazia, as these plants differ from each other with morphological features and with separated areas. These plants differ from each other so distinctly, that acknowledging P. Wittmanniana Hartwiss, as a species, A. A. Grossheim joined the Talishsky peony with P. Mlokosewitschii Lomak. from the Lagodekhsky gorge, that was quite incorrect. And if any joining of this kind was to be done, it would be better to follow the example by Stern, considering this plant a synonym P. Wittmanniana Hartwiss ex Lindl. (non Stev.). Nevertheless, the Talishsky yellow-flowered peony is a self-dependent species and the name P. tomentosa (Lomak.) N. Busch should be preserved for it.

From all Caucasian red-flowered peonies Stern recognized only two species P. daurica Andrews and P. tenuifolia L. and regards P. caucasica N. Schipcz. to be a synonym of the former species.

P. daurica Andrews - P. triternata Pall. has been described in the Crimea and is characterized with rounded, at both sides blue-gray leaf-lobes, with pink stigmas and pink tomentose pubescence of ovaries.

I have never encountered peonies with the above-indicated characters in Caucasus and such the peonies are not to be taken for the Caucasian red-flowered peonies. The latter peonies are not homogeneous, so that, besides P. caucasica N. Schipcz, two new species have been discovered by us, which are P. Ruprechtiana Kem.-Nath. from the regions bordering with Imeretia, Kartii and Meskhetia and P. lagodechiana Kem.-Nath. from the Lagodekhsky gorge. Neither P. coralliana Retz., nor P. Kavachensis grows in Caucasus; the former south-european and mediterranean species with narrow lance-shaped leaf-lobes is well-distinguished from the Caucasian species, and yet in 1903 N. A. Busch denied its existence in Caucasus.

As to P. Kavachensis Aznaur. described in the Minor Asia and presented by A. A. Grossheim for the Caucasus Flora, Stern is' quite right separating it from P. caucasica and presenting it, as a synonym of P. Mascula Mill.; and at the same time P. corallina Retz. and P. caucasica N. Schipcz. are identified by him with P. daurica Andrews, that is not correct.

We don't support the idea of enlargement species in such a way like this. Theoretically, it is possible to decrease a number of Caucasian peonies species to the further extent, to three, e.g., recognizing P. Wittmanniana Hartwiss ex Lindl. with yellow blossoms, P. daurica Andrews with red blossoms and P. tenuifolia L. with leaves manifold dissected into narrow lacinules. It seems so simple to do this.

However, the species defined in the above-mentioned way would be formal, "speculative", but not concrete real species; "in most cases systematizers-monographers and florists deal with geographical races, of which some of them may correspond to small species." ... "a species is important for us, as a homogeneous population occupying a concrete place within the frame-work of economic relations in Nature, and it does not even matter in what way this species has arisen", (the academician V. L. Komarov, 1940).

Only, taking into consideration such an approach like this, it is possible to get along with comprehending the Caucasian peonies on the whole and the species from the section Sternia Kem.-Nath. in particular, (about species from two sections Flavonia and Paeon, it was already mentioned above).

From this sections for Caucasus, Stern presents only P. tenuifolia L. which was described in Ukrainian steppes and from where it propagates reaching Hungarian steppes. But as a matter of fact, there are far more of these peonies in Nature, as it was indicated by some old authors. E.g. Ruprecht has described P. Biebersteiniana Rupr. found upon the Stavropolsky hills and presented it in-Flora USSR, as a self-dependent species; besides, quite recently two new species from Kartly have been described by the academician N. N. Ketzchoveli and which are as follows: P. carthalinica Ketzch. for which a brief description has been presented in Georgian, in 1937 (submitted by us in Flora Georgia, in 1948) and P. Majko Ketzch which has been described in 1959.

The former species is very closely related with the Stavropolsky peony. Besides, along with P. tenuifolia L they form the evolutional series Ftssae Kom., established by the academician Komarov in 1937. The latter species P. majko Ketzch. has been firstly found upon a margin of a light forest, upon hills, among thickets formed by Paliurus spina christi, in 1939; it well differs from the representatives of the series Fissae Kom. with its broad leaf-lobes of pinnately dissected leaves and with its large pink flowers, being closely related with the Mediterranean species P. hybrida Pall. and the Chinese P. Veitchi Lynch, for which the new series Hybride Kem.-Nath. has been established by us.

The kindred relations between Caucasian peonies based on their morphological characters and distribution areas may be represented in the way, as follows.

A territory of the Ancient Katasia and Caucasus is presumed by us to be Motherland of the genus Paeonia L. Hence, the contemporary Caucasian species have the autochton origin.

Being more ancient, the species from the section Flavonia are as follows: P. macrophylla (N. Alb.) Lomak. and P. Steveniana Kem.-Nath. They are closely related with Chinese species from the serieses Luteae and Lactiflorae; from them. P. macrophylla is the most ancient species in accordance with its morphological characters (naked ovaries and fruits, very large, broad, green at both sides leaf-lobes), ecology and area of distribution (Adjaria, which is the most humid part of Caucasus).

Its ancestors had been presumably propagated all over Caucasus, in montane forests of the tretiary period; but as environmental conditions had been changing towards aridity, the ancestral species had begun acquiring new features, such as pubescence upon the inferior side of leaves and more deep colour of petals, and afterwards, tomentose pubescence of ovaries and fruits and, at last, blue-gray bloom upon leaves.

Afterwards, the ancestral species had splitted into species, from which P. macrophylla (N. Alb.) Lomak. populated Adjaria; being very closely related with P. macrophylla, P.

Steveniana Kem.-Nath. (P. Wittmanniana Stev.) propagated within the Borjom-Bakuriansky province, Meschetia; in Abchazia, P. Wittmanniana Hartwiss ex Lindl. had arisen which had tomentose-downy ovaries and fruits and green at both sides leaves; in the mountains of Talish, P. tomentose (Lomak.) N. Busch had got formed, as species, having tomentose-downy ovaries and fruits; and at last, in the mountains of Lagodechy and adjoining forests of Dagestan, P. Mlokosewitschii Lomak., a species with tomentose-downy ovaries and fruits, pink stigmas and glaucescent leaves, had settled separately from others.

Simultaneously with peonies from the section Flavonia Kem.-Nath., the ancestor of Caucasian representatives from the section Paeon DC. changed also. It had splitted into the species, as follows: P. Ruprechtiana Kem.-Nath. which grew in montane forests at the border between Imeretia and Kartly and was characterized with naked, green at both sides, lustrous leaf-lobes and pink blossoms; (we consider it more closely related with the ancestral species which had been growing all over Caucasus); in Lagodechsky gorge and forests of Dagestan, P. lagodechiana Kem.-Nath. with pink flowers and tender leaves had arisen; and, at last, P. caucasica N. Schipcz. with red or purple flowers and with leaves, which were glaucescent at one side only, propagated all over more spacious area.

As to Caucasian peonies from the section Sternia Kem.-Nath. which grow in steppes and turned-into-steppe places of the North Caucasus (Predcaucasiye) and TransCaucasus, they have more ancient origin. As it is well-known, in Caucasus, during the quaternary period, such the environmental conditions had been coming to existence, which were necessary for forming the steppe flora. Its specific composition might have been consisted of elements of different origin, such as the Caucasian montane-xerophylous element, firstly, the caucasian-forest element, secondly, and the migration element, lastly.

There are many reasons to suggest, that our forest peonies from the section Paeon DC. might have been taking part in forming the steppe flora of Caucasus. From them some special form with narrow dissected leaf-lobes might have developed and populated forest margins bordering with steppe and afterwards, propagated all over the North caucuses (Predcaucasiye) steppes and far to the North, reaching the South-Russian and West-European steppe regions.

Also, a possibility exists to accept their migrational origin. From Siberian steppes along the north and south coasts of the Caspian Sea, the mutual ancestor of our Caucasian peonies and P. hybrida Pall. could have propagated, from which the youngest descendant is to be considered P. tenuifolia L. in this case. The kindred relations for our Caucasian peonies may be represented by the Scheme 2 .

It should be emphasized that studying a history of the Caucasian flora with respect to the genus Paeonia L. is quite important because there are the genus Paeonia L.'s representatives, especially, yellow-flowered peonies, growing in the forests of TransCaucasus and because these Caucasian peonies occur to be closely related with the chinese-himalayan yellow-flowered peonies, forming with them one mutual section. All this means that the transcaucasian forests represent a remnant of the ancient flora which is more or less preserved in the South-East Asia nowadays and which might have been including the Caucasian, South European and maybe even North African flora quite too long ago, in the tretiary period or maybe even earlier. In this case the Caucasian peonies, especially the representatives of the section Flavonia Kem.-Nath., happen to be an echo of this remote epoch. The above-mentioned section may serve as an indication to one of the particular cases of floristic relations between Caucasus and the East Asia.