F.C. Stern A Study of the Genus Paeonia

chapter IV


part 4 1901-

1901 N. A. Busch in Kusnezow, Busch, and Fomin, Flora Caucasia critica, 3, iii, 14, describes the different species found in the Caucasus region. P.tenuifolia with its habitat in the Crimea and the steppes of the Caucasus and Transcaucasia grows at about 3000 ft. ; then a variety Biebersteiniana (Ruprecht) N. Busch with broader segments than P.tenuifolia is described, giving as a synonym P.hybrida of Pallas, which is also found in the same environment. P.triternata {P.daurica) is named [end page 138] P.corallina subsP.triternata and recorded from all the Caucasus at 3000-6000 feet as well as the Crimea, and a variety coriifolia (Ruprecht) N. Busch from Transcaucasia and differing from P.triternata by the hairs on the underside of the leaves.

This publication is particularly important for the full details given of the habitat of each of the yellow-flowered species. P.Wittmanniana of Steven, which is P.Wittmanniana var. nudicarpa with glabrous carpels, comes from Abkhazia in western Transcaucasia, and also the forma macrophylla with large leaves from the same district. The subspecies tomentosa (now named P.Wittmanniana Lindley), only separated, as he says, from P.Wittmanniana by the densely tomentose carpels, comes from the district of Talysch which borders the Caspian Sea. Busch adds that Buhse found a yellow paeony at Massula in Persia ; the range of mountains in the Talysch district continues on to Massula in Persia. Bunge reports that he found P.Wittmanniana near Asterabad in Persia. This is probably P.Wittmanniana (syn. P.Wittmanniana var. tomentosa). Asterabad is situated at the south-eastern comer of the Caspian Sea just at the east end of the Elburz Mountains; P.Wittmanniana has been found in the Elburz Mountains. The habitat of P.Mlokosewitschi is given as in the woods near Kachetia, which is about the centre of the Caucasian range.

In the Addenda to this volume, dated 1903, there is a note that since the original was published, material of P.Wittmanniana with tomentose carpels has come from Abkhazia, the district near the Black Sea. This discovery has further complicated the difficult problem of the exact location of these yellow-flowered paeonies.

1901 Paeonia lutea is described and illustrated in the Botanical Magazine, 127, t. 7788. Sir Joseph Hooker recounts how this interesting plant has been received from Professor Cornu from the Jardin des Plantes, Paris, and how it had flowered in June 1900 in the Temperate House at Kew.

1903 In the second volume of the Icones ad Floram Europae, by Jordan and Fourreau, six paeonies are described by Jordan with entirely new names ; there are coloured illustrations of them. The names are as follows and their correct designations, as far as I can judge, should be :—


Hautes Alpes

.. P.officinalis.


.. P.officinalis.



.. P.humilis.


.. P.humilis.



.. P.Russi.


.. P.Russi var. Reverchoni.

1904 Fedtschenko in Fl. Thian Shan in Acta Horti PetroP.23, 353, writes, " in 1830 C. A. Meyer (in Ledebour Flora Altaica, 2,277) described the two species, namely P.hybrida Pallas and P.intermedia. In 1860 Trautvetter reunited both with P.anomala and distinguished a typical form with glabrous carpels and a variety hybrida with hairy carpels, and as a subform of the latter a forma intermedia (C. A. Mey.) with broader segments of the leaves. In 1901 P.N. Kryloff in Flora Altaica described from his region two species, namely P.anomala L. and P.hybrida Pall. In my opinion P.anomala L. sens. lat. does indeed represent the two subspecies of which the one with hairy carpels is met within our area. For this subspecies I retain the name intermedia proposed by Meyer, because the name hybrida has been applied to different forms and indefinitely and because it is rather doubtful what Pallas did really understand by this name."[end page 139]

1907 Paeonia Cambessedesii is described and figured in the Botanical Magazine, 133, t. 8161. The illustration is poor but it does bring out two of the unique characters of this species—the deep red back of the leaves and the large number of carpels. Dr. John Hutchinson describes the species and says that Miss Frances Geoghegan of GIasnevin, Dublin, introduced it to cultivation.

1908 Paeonia Mlokosewitschi is illustrated in the Botanical Magazine, 134, t. 8173. The late Dr. Stapf discusses the yellow-flowered species of the Caucasus, quoting Steven, Lomakin and Alboff. He says this species was discovered by Mlokosewitsch near Lagodekhi in the eastern part of the central Caucasus.

1909 P.Veitchii is first described by R. Irwin Lynch of the Botanic Gardens, Cambridge University, in the Gardeners' Chronicle, 3rd series, 46, page 2. He shows the difference between this species and P.anomala and stresses the character of P.Veitchii having more than one flower to the stem.

1910 In the Gardeners' Chronicle, 48, page 366, there is an article describing P.japonica (Makino) Miyabe and Takeda, by Professor H. Takeda, who says that this paeony has been confounded with P.obovata, which also grows in the mountainous districts of Japan, but P.japonica is separated from Maximowicz's plant by the flowers which open less widely, the more concave petals, short stigmas and glabrous leaves. Makino described this Japanese Paeony as P.obovata var. japonica in the Botanical Magazine, Tokio, 16, 59 (1902).

1910 J.Briquet in his Prodrome de la Flore Corse recognises two species indigenous to Corsica, making them varieties of P.corallina ; P.corallina var. pubescens Moris. (1837) and P.corallina var. leiocarpa Coss. (1850) with glabrous carpels, from Monte Cagna among other places. The first-named would be P.Russi and the second P.Russi var. leiocarpa.

1915 The first description of P.Mairei, which was found by E. E. Maire in Yunnan in 1911 and 1912, was published by Leveille in Bulletin de I'Academic Internationale de Geographie botanique, 24. He describes the carpels as densely tomentose with golden hairs.

1916 In Flora Cretica, by Michael Gandoger, there is a list of the flora of Crete with the habitats of each species, including P.corallina and P.cretica.

1916 Paeonia Willmottiae is illustrated in the Botanical Magazine, 142, t. 8667, and described as a new species by Dr. 0. Stapf. He says that this species approaches P.obovata but can be distinguished from it by the larger leaflets, glaucous and tomentose below, by the white flowers and by the pistils which considerably exceed the relatively short anthers. The plant from which the excellent illustration was drawn appeared in a pan of Chinese seed collected by E. H. Wilson and raised by Miss Willmott at Warley Place, Essex.

1917 G.V.Aznavour in Magyar Botanikai Lapok, 16, 7, describes as a new species P.kavachensis from Vastan in the district of Kavache, Armenia. From the description this paeony would appear to be P.mascula. It is interesting that Aznavour notes that there is a monastery of St. Thomas in this district.

1918 In the Botanical Magazine, 144, t. 8742, there is a beautiful illustration of Paeonia peregrina Mill. and a most interesting discussion by Dr. 0. Stapf how this name " peregrina " came to be confused between the Balkan paeony and a paeony from the south of France ; the confusion is fully discussed on page 127.[end page 140]

1920 and 1921 There are several mentions of paeonies in Notulae systematicae ex Herbaria Horti Botanici Petropolitani of 1920 and 1921. In no. 7, 1920, P.bifurcata is described by N. Schipczinsky as a new species. The specimen examined was no. 566 of Pere Farges from Tchen-keou-tin, north Szechwan. This paeony has been included in this study under P.Mairei. In vol. 2, 1921, V. L. Komarov describes two new species as P.Beresowskii and P.Potanini. P.Beresowskii has been found to be the same as P.Veitchii. P.Potanini has been adopted as the specific name of the small flowered species in the subsection Delavayanae.

N. V. Schipczinsky has a brief survey of the genus in 11, 1921, with a key mainly following Huth, and a list of species dividing some of them into varieties and giving their habitats. Under P.Wittmanniana Lindl. he has two varieties, var. nudicarpa with glabrous carpels and var. tomentosa Lomak. with tomentose carpels. Under P.corallina he has a variety caucasica which would appear from the description to be P.daurica.

1930 In Flora Kavkaza, II, 90-92, by A. A. Grossheim, there is a key to the paeonies of the Caucasus. P.Biebersteiniana of Ruprecht is made a variety of P.tenuifolia. This paeony is described with broader leaf-segments and very short bristles along the veins on the upper side of the leaf. This reads like the description of P.anomala var. intermedia. P.corallina var. caucasica might from the description be P.daurica ; the leaves are described as glabrous or pubescent below ; the leaves of P.daurica are usually glabrous, though there are plants in gardens received from the Tints Botanical Garden which are without doubt P.daurica, that have a pubescence of short hairs on the underside of the leaves. The leaves of P.corallina var. coriifolia are described as hairy below, otherwise the description is the same as the last.

The yellow-flowered paeonies are fully described. The paeony with glabrous carpels is named P.Wittmanniana Stev., and the large-leaved form is named var. macrofthylla. The paeony with tomentose carpels from the Lenkoran district is named P.tomentosa (Lorn.) N. Busch, with another paeony with tomentose carpels from Abkhazia described as P.abchasisa (sic) Mischenko with the leaflets glabrous or not densely hairy on the veins beneath. This being one of the two differences between these paeonies with tomentose carpels; the other difference is that P.abchasisa has leaflets 17-18 cm. long by 10 cm. broad compared to the leaflets of P.tomentosa which are 6-8 cm. long by 3-6 cm. broad.

1931 Handel-Mazzetti gives a full description in Symbolae Sinicae, 7, 265, of P.oxypetala, a new species found by him in the province of Szechwan in western China and first described by him in 1920 ; he says that this species is unique in the whole genus by the petals having the apex acute.

1931 In the Botanical Magazine, 155, t. 9249, there is a plate of Paeonia tomentosa with a description and history of the plant by the late Dr. 0. Stapf. Dr. Stapf says the only obvious difference between this plant and P.Wittmanniana Stev., is the presence of a dense tomentum on the carpels of P.tomentosa and its complete absence in the other. The form with tomentose carpels is now known as P.Wittmanniana Lindl. and the one with glabrous carpels as P.Wittmanniana var. nudicarpa. Dr. Stapf then enumerates four pairs of paeonies which are almost solely separated by the presence or absence of tomentum on the carpels. He has an interesting argument on this phenomenon saying that these types are highly stable as no intermediate states occur between them even in cultivation, where most of them have been for a long time without showing any abrupt or gradual change from one type to the other ; there is a sharp distinction between the species with glabrous and tomentose carpels running through the genus, though not always resulting in the appearance of parallel forms like those enumerated. Therefore he argues these types, which are mutually [end page 141] exclusive over large areas and may therefore be considered as genetically fixed within those areas, should be accepted as specifically distinct. (See page 9.)

1937 In Flora U.R.S.S. 7, the species of the genus Paeonia found in Russia are dealt with by Schipczinsky and Komarov. There is a key and the text is in Russian. There is much interesting information about the yellow-flowered paeonies of the Caucasus ; this information is incorporated in the text of the descriptions of P.Wittmanniana and the two varieties, and P.Mlokosewitschi. P.caucasica is included as one of the species found in the Caucasus. The synonyms given are P.corallina var. caucasica N. Schipcz., P.corallina subsP.triternata Busch, pro parte, P.triternata Ruprecht (1869). There is also a drawing of a leaf of this species which might be P.daurica. No mention appears to be made of the hairs on the underside of the leaflets of P.caucasica in the key.

1938 Professor G. Ledyard Stebbins, jun., has written several important papers on the genus. The Cytogenetic studies (1938) have already been referred to (page 12). In the University of California publications in Botany, 19, no. 7, 245-266 (1939) there are Notes on some systematic relationships in the genus Paeonia by Stebbins. In this paper which has been referred to elsewhere in this study, the morphological criteria of the species is discussed and the character of the sepals is emphasized. There is a key to the species belonging to the subgenus Paeon. It is not always quite clear whether the names of the species given by Stebbins are the same plants as referred to under those names in this study. There is much interesting and useful information of the crossing of the different species by Professor A. P.Saunders. The American Subsection Onaepia has been dealt with by Stebbins in an exhaustive and valuable paper in Madrono (1938), 4, no. 8, 252-260. This paper has been fully referred to under the species P.Brownii in this study.

1939 In Acta Horti Gotoburgensis, 13,37-40, the late Dr. Heinrich Handel-Mazzetti has an important paper on the paeony species to be found in China with details of their distribution. The localities for P.lactiflora, P.Veitchii and P.lutea are given without comment. P.anomala is recorded from the province of Shansi and he also mentions P.Veitchii var. Beresowskii (Komarov) N. Schipcz., from north Szechwan, and adds a note after the latter that one specimen has almost glabrous carpels and one has quite glabrous carpels ; further, that single-flowered specimens are found and it is not clear how these differ from P.anomala. Probably all these specimens are P.Veitchii ; a full account of the difference between P.Veitchii and P.anomala is given under the description of P.Veitchii.

A Latin description of P.Mairei is given with several new locations ; the carpels are described as covered with vesicular papillae, and P.oxypetala only differs from it by the acute petal and the hairy covering of the carpels. The locations of P.suffruticosa are given as temple gardens but a new shrubby species P.decomposita Hand.-Mzt. is described. I have seen a photograph of the type specimen (H. Smith, no. 4641) of P.decomposita from the Herbarium of the Botanical Museum, Upsala, and if this photograph is compared with Rock's specimen of P.suffruticosa in the Kew Herbarium they will be found to be alike.

Handel-Mazzetti also refers to P.obovata var. glabra Mak. in Journal JaP.Bot. 5, 35 (1928), specimens with glabrous leaves from Shansi, Chihii, Manchuria and north-east Yunnan.

1940 In the Botanical Magazine, 162, t. 9594, is recorded the change of name of the Cretan paeony —P.cretica of Clusius—to Paeonia Clusii. The necessity for this change of name is fully set out under the description of P.Clusii. [end page 142]