Paeonia intermedia(1)
Herbs perennial, up to 70 cm tall. Tap roots cylindrical, to 2 cm in diameter, woody when old; lateral roots thickened, tuberous, tubers spheroidal to long-fusiform. Lower leaves biternate, covered with bristles along veins above, always glabrous beneath; leaflets several times segmented; segments 70—100 in number, more-or-less decurrent at the base, linear, 6-16 cm long, 0.4-1.8 cm wide, sometimes lobed, acuminate at the apex. Flowers solitary, terminal; involucrate bracts 3 in number, leaf-like, unequal in size; sepals 3-5 in number, often red-purple, ovateorbicular, 1.5-2.5 cm long, 1-2 cm wide, mostly rounded (at least 2 non—caudate) at the apex, glabrous; petals 7—9 in number, purple-red, obovate, 3.5—5.5 cm long, 1.5—3 cm wide, irregularly incised at the apex; filaments purple; anthers yellow; disk annular, incised, up to 2.5 mm high; carpels 2—5, but mostly 3 in number, tomentose, rarely glabrous; stigmas sessile, 1 mm wide, red; ovules 12-16 per carpel. Follicles 2—2.8 cm long, 1.1—1.3 cm wide. Seeds black, glossy, long-ovoid, 5-5.5 mm long, 3-3.5 mm in diameter.

P. hybrida – Copyright Andrey Dedov

Chromosome number: 2n=10 (diploid).

Growing on grassy and shrubby slopes, in meadows, steppes, or sparse woods, at altitudes from 900 to 3,250 m. Widely distributed in N Xinjiang of China (S to the Tianshan), Kazakhstan, Kyrghyzstan, Tajikistan, Uzbekistan, and the Altai of Russia.

Until Hong & Pan (2004), Paeonia intermedia had long been identified as P. hybrida (Krylov, 1901; Schipczinsky, 1921, 1937; Gamaulova, 1961), or treated as a variety of P. anomala (Fedtschenko & Fedtschenko, 1905; Stern, 1946; Pan, 1979), as a subspecies of P. anomala (Trautvetter, 1904), or even as a form of P. anomala var. hybrida (Trautvetter, 1860). These different and erroneous assignments were made because: (1) the identities of P. anomala, P. intermedia and P. hybrida were not clear to the authors mentioned above, (2) previous authors emphasised the taxonomic value of indumentum on carpels, and (3) the root and calyx characters and their correlation were ignored. Our examination of the types of these three taxa, together with extensive observations of herbarium specimens and natural populations, show that P. intermedia C. A. Mey. is an independent species, differing distinctly from P. anomala by its lateral tuberous to long—fusiform roots, and sepals that are mostly (at least 2 out of 3-5) rounded at the apex but not caudate (Hong & Pan, 2004). In contrast to P. anomala, P. intermedia prefers relatively sunny and dry habitats. In P. intermedia carpels vary in number from 2 to 5, mostly 3, and from tomentose to rarely glabrous even within populations. Ovczinnikov (1975) properly treated the peony in Tajikistan as an element of P. intermedia, but his description of the new subspecies pamiroalaica is not justifiable. His description and our extensive observations have not revealed any significant difference from other populations. Paeonia hybrida Pall. has been shown to be a synonym of P. tenuifolia L. (Hong & Pan, 2004).

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Editor’s note:

After some discussion with people that have collected wild peonies in Altai and Central Asia, we have added another subdivision of this species:
-Roots fusiform: subspecies intermedia
-Roots tuberous: subspecies hybrida

Remark that the more logical classification would be Paeonia hybrida ssp hybrida and Paeonia hybrida ssp intermedia, but we currently keep the former classification to keep it in line with Hong’s classification and key to the species.

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Editor’s note:
Paeonia hybrida from the Altai region in Russia is by many people considered to be an independent species or a (natural) hybrid between P. tenuifolia and some other species and thus the treatment by Hong of P. hybrida as a synonym for P. tenuifolia is contentious. We however, consider it to be best treated as a subspecies in P. intermedia. We would like to bring forward the following reasons for this:

  • The first description of P. hybrida was by Pallas in 1789, but “Pallas (1789) described … P. hybrida from a plant raised from seeds of P. tenuifolia cultivated in the Botanic Garden of the St. Petersburg Academy in Russia.”(2) Thus this plant, the so-called type-specimen, is at least 50% P. tenuifolia and perhaps even 100%. It is obvious that Hong would come to the conclusion that P. hybrida Pall is a synonym for P. tenuifolia. The type-specimen is simply not representative for what most people consider to be P. hybrida, which is supposed to grow wild in the Altai mountain steppe of Russia.
  • The “P. hybrida” (a population of only 5 plants) occurring in Georgia is not a separate species but a natural hybrid between P. tenuifolia and P. caucasica (=P. daurica coriifolia). It is known locally as P. majko and has nothing to do with either P. anomala, P. hybrida or P. intermedia.(3)
  • The hybrid plants that derive from crossing P. tenuifolia with P. anomala are sometimes called P. hybrida, but do not occur naturally and are thus ‘garden hybrids’. The garden hybrids “P. hybrida” usually have much narrower leaflets in larger numbers than P. hybrida from the Altai. It is unwise and confusing to refer to these garden hybrids as “P. hybrida” as they have nothing to do with the peonies in the Altai where P. tenuifolia does not even occur.
  • The roots of P. anomala are carrot-shaped, whilst the roots of P. intermedia and P. hybrida are either fusiform-tuberous or tuberous. The latter two are thus clearly distinct from P. anomala but are hard to discern one from the other as is also confirmed by DNA-analysis by Efimov et al.: “In comparison to morphological data, molecular data allow to differentiate more accurately these species, especially to distinguish P. anomala from P. intermedia and P. hybrida.”(4) The data from this research concludes that P. intermedia is very close to P. hybrida genetically and it is unsure whether distinct enough to separate them as the same species in different locations or distinct species themselves. The morphological differences are twofold according to this publication. First are the roots: P. anomala has carrot-shaped roots, P. hybrida has tuberous roots and P. intermedia falls in between with fusiform thickened roots. Second is the habitat: P. anomala is a mesophyte (growing under medium conditions of moisture), growing in forests, forest edges, glades; P. hybrida at the other extreme is a xerophyte (growing under dry conditions) common in the steppe region; P. intermedia lives in mountains, foothills, and mountain-steppe forests. The three species all grow in the Altai, P. intermedia also grows more southernly in Central Asia and P. anomola has a very wide northernly distribution. It is as yet unclear whether P. hybrida also grows outside of the Altai. DNA research is able to distinguish P. hybrida (Altai) from P. intermedia growing in Central Asia, but not from P. intermedia growing in the Altai itself.
  • The most obvious difference between P. hybrida and P. intermedia would be the form of the roots with P. hybrida having tuberous roots, resembling those of P. tenuifolia, and P. intermedia having fusiform tuberous roots with the thickened part of the roots further away from the crown. However, Hong visited the natural populations and concluded this: “Gamaulova (1961) described roots of Paeonia anomala as fusiform-thickened, and those of P. hybrida as tuberous-thickened. However, roots of P. intermedia vary in shape from tuberous consistently to fusiform, and this variation could be found within populations or even on the same individual, whereas in P. anomala neither fusiform-thickening nor tuberous-thickening were found.”(5) It is thus clear that this character can help to distinguish between P. intermedia/P. hybrida on the one hand and P. anomala on the other. But clearly not between P. intermedia and P. hybrida. It seems Gamaulova misinterpreted P. intermedia for P. anomala, which then resulted in further confusion in future publications and the discovery of distinct ‘new’ species with carrot-shaped roots like P. sinjiangensis and P. altaica which are in fact nothing but synonyms for P. anomala.
  • Other sometimes mentioned differences are either continuous characters (presence/absence of indumentum on the carpels) that can all be found in one population or may be due to habitat differences (height). The only difference we are unsure of is in the seeds, which should be black in P. intermedia and brown in P. hybrida, although this may be due to the maturity of the seeds.(6)
  • The findings of Hong are at odds with Efimov. According to Hong the different root systems can be found in the same populations and even plants. It has to be said however that only populations in the Altai were sampled and not P. intermedia populations further South in Central Asia. This solves the difference with Russian research by Efimov that was able to discern P. hybrida from P. intermedia in Central Asia, but not in the Altai itself. Efimov considers the root systems of P. intermedia to be intermediate between P. hybrida and P. anomala. The habitat shows the same differences. We currently conclude that P. hybrida is probably best regarded as a subspecies for P. intermedia, which may in due time become a full species.

Images below by Andrey Dedov
Paeonia intermedia ssp hybrida (or Paeonia hybrida var. typica) in steppe grasslands, Altai.
“P. hybrida var typica grows in the Altai steppe in the area, has roots round brown seeds. height 25-35 cm.”

Footnotes:
  1. Hong, De-Yuan. “Peonies of the World. Taxonomy and phytogeography.” Kew: Royal Botanic Gardens, 2010, p. 205[]
  2. Hong, DY & Pan, KY. (2004). A taxonomic revision of the Paeonia anomala complex (Paeoniaceae). Annals of the Missouri Botanical Garden. 91. 87-98 []
  3. Punina EO, Machs EM, Krapivskaia EE, Kim ES, Mordak EV, Miakoshina IuA, Rodionov AV. “Interspecific hybridization in the genus Paeonia (Paeoniaceae): polymorphic sites in transcribed spacers of the 45S rRNA genes as indicators of natural and artificial peony hybrids”. Genetika. 2012 Jul;48(7):812-26.[]
  4. S. V. Efimov, G. V. Degtjareva, E. I. Terenteva, T. Kh. Samigullin, C. M. Valiejo-Roman. “Phylogenetic relatioships of the Paeonia anomala, P. intermedia and P. hybrida (Paeoniaceae) inferred from nuclear ITS rDNA and plastid ycfl gene.” In: Проблемы ботаники Южной Сибири и Монголии» – XV Международная научно-практическая конференция, 2016, pp. 112-116.[]
  5. Hong, DY & Pan, KY. (2004). A taxonomic revision of the Paeonia anomala complex (Paeoniaceae). Annals of the Missouri Botanical Garden. 91. 87-98[]
  6. Description of both species by Andrey Dedov[]

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