The genus Dionysia (Primulaceae), a synopsis and five new species

Willdenowia 37 – 2007 37 

MAGNUS LIDÉN 

The genus Dionysia (Primulaceae), a synopsis and five new species 

Abstract 

Lidén, M.: The genus Dionysia (Primulaceae), a synopsis and five new species. – Willdenowia 37: 

37-61. – ISSN 0511-9618; © 2007 BGBM Berlin-Dahlem. 

doi:10.3372/wi.37.37102 (available via http://dx.doi.org/) 

Five new species of Dionysia are described (D. viva, D. zschummelii, D. crista-galli, D. zetterlundii 

and D. tacamahaca) from the Zagros mountains of W Iran. One subspecies (D. sarvestanica subsp. 

spatulata), one variety (D. gaubae var. macrantha) and two sections (D. sect. Zoroasteranthos and D. 

sect. Mucida) are also described as new. A revised classification of the genus is suggested and a 

complete list of species is provided, in which some emendations to previously published data are 

given. A dichotomous key to the species is presented. 

Key words: Iran, Zagros Mts, Primula, cushion primroses, taxonomy. 

Introduction 

The genus Dionysia Fenzl, belonging in Primula sensu lato, is a typically Irano-Turanian genus 

almost restricted to the rather dry mountains of the Flora Iranica area, from SE Anatolia and W 

Iran to Tadzhikistan and Afghanistan. Most species are cushion forming chasmophytes – in spring 

completely smothered in yellow, purple, violet or pink flowers – but some are more lax and 

“Primula looking”. Some species are very choosy about their habitat and only grow below overhangs, 

whereas other species can be found also on slightly sloping rocks. In spite of being difficult 

to grow, a large number of species are cultivated by devoted specialists or in botanical gardens. 

When Per Wendelbo (1961a) revised the genus Dionysia, he recognised 28 species, but 

added several more in later publications. 41 species were known to science when Christopher 

Grey-Wilson (1989) published his semi-popular account “The genus Dionysia”, in which all species 

are described and illustrations (including several colour photographs) and distribution maps 

are provided. The number of recognised species in the present paper is 49 and there is every reason 

to suspect that there are even more species waiting to be discovered, as several are known 

from a single locality and/or based on very recent finds. 

The number of species in cultivation has also increased dramatically in recent years, from a 

handful in the 1960s, to an estimated 21 in 1989, when Grey-Wilson wrote his book. Today 44

38 Lidén: Dionysia, synopsis and new species 

species are grown, representing 90 % of the known taxa. As live plants are indispensable for a 

proper understanding of morphology and biology, plants in cultivation have contributed considerably 

to current progress in our knowledge of the “cushion primroses”. 

Molecular investigations by Ida Trift & al. (2004) have given us a far better understanding of 

evolution and biogeography of Dionysia. Recent field expeditions have brought us important observations 

on their habitat and many informative photographs of flowering individuals in the wild. 

Several species are considered threatened by nature conservation authorities in Iran, mainly 

due to their restricted distributions, but some populations are also subjected to collection for 

medicine. Because of their beauty, rarity and dramatic stations, Dionysias are among the few 

plants referred to in Iranian tourist information. 

Material and methods 

This paper is based on experience from field studies in Iran (Zagros), plants cultivated in greenhouses 

at Göteborg Botanic Garden and herbarium studies. It takes as a starting point the monograph 

by Per Wendelbo (1961, with emendations up to 1980), which is accepted as to species 

level taxonomy. Types (at least isotypes) and other cited herbarium specimens have been seen unless 

otherwise indicated. 

Since Wendelbo’s monograph, rich herbarium material has accumulated in the herbarium of 

the Institute of Forests and Rangelands (TARI) in Tehran. During the same period, hardly anything 

has reached other major herbaria. In addition, some recent expeditions to Iran, notably 

SLIZE (Swedish-Latvian-Iranian-Zagros-Expedition, with Assadi, Mozaffarian, Popp, Seisums 

& Lidén) in 1998 and T4Z (Tjerdsma, Zetterlund, Zschummel & Zschummel to Zagros) in 2002, 

have brought more than 60 accessions of 31 taxa into cultivation. This means that I have had the 

opportunity to study a far richer material from Zagros than Wendelbo, Grey-Wilson or even 

Jamzad had available. Of the Iranian species D. bornmuelleri, D. tacamahaca and D. sawyeri has 

not been studied in cultivation. 

The following herbaria were consulted, directly or indirectly: A, B, BASBG, BG, BR, C, 

DBN, E, GB, K, LD, LE, MB, NY, P, PE, PR, PRC, S, Shiraz University, TARI, W (abbreviations 

according to Holmgren & Holmgren 1998-). 

Unless otherwise indicated, single measurements of leaves, calyx, etc. always refer to length. 

Taxonomic history 

Dionysia was first recognised as a genus by Fenzl (1843), based on material of D. odora collected 

by Kotschy in Kurdistan. This, however, was not the first Dionysia known to science. Already 

in 1817 Lehmann had described Primula aretioides on material collected in N Iran in 

1770, but only in 1846 Boissier formally transferred it to Dionysia, at the same time describing 

three new species. In the meantime Duby (1844) had treated the Primulaceae for Candolle’s 

Prodromus, in which he included the Dionysia species in Gregoria, a synonym of Douglasia, a 

rather remotely related genus. To complicate matters, Duby later the same year erected the genus 

Macrosyphonia, to which he transferred his Gregoria caespitosa. 

Bunge (1871) published an account of Dionysia, recognising it as a genus separate from 

Primula, as did Boissier (1879). Kuntze (1891) merged Dionysia into Primula as a section. 

Knuth (1905) treated Dionysia for Engler’s Pflanzenreich, recognising 20 species. Bornmüller 

(1899, 1903, 1904, 1905, 1910, 1937; for references, see the List of species, below) was ambiguous 

concerning generic status; although he accepted Dionysia as a genus, he was careful to ensure 

that his new names should remain valid also in Primula, and even provided the alternative 

combinations, in one case a nomen novum. 

Melchior (1943) discussed the evolution and subdivision of Dionysia. His thoughts were largely 

followed by Wendelbo (1961), whose exhaustive monograph has been the basis for subsequent 

revisions, of which the most important are Grey-Wilson (1989), with a forerunner (1970),


and an account of Iranian taxa by Jamzad (1996). All in all, Grey-Wilson contributed three and 

Jamzad four new species. 

An evolutionary approach to the classification of Dionysia has now been made possible by 

the recent molecular studies by Mast & al. (2001) and Trift & al. (2004). These authors present a 

phylogeny, which in parts differs drastically from the speculative schemes of Melchior (1943) 

and Wendelbo (1961a), which involved now obsolete concepts like “advancement level”. 

Morphology and relationships 

That Dionysia, though surely a natural group in itself, can not be demarcated from Primula has 

been obvious from the start. Both Wendelbo (1961a) and Grey-Wilson (1989) realised that Dionysia 

belongs in Primula in a genealogical classification, but as both accepted a polyphyletic 

Primula, they saw no problems in recognising Dionysia as a genus, following the tradition of 

Bornmüller and Melchior. 

Dionysia differs from most Primula in the base chromosome number 10, the suffruticose 

habit, long corolla tube and a capsule that splits to the base into 5 valves. Further, several species 

are characterised by woolly farina, which is very rare in Primula (Wendelbo 1961a) but present in 

P. qinghaiensis (Trift & al. 2002). Efarinose species and species with powdery farina are also 

known in Dionysia. All characters distinguishing Dionysia can be found in other groups of Primula, 

though never in combination, and accordingly Dionysia can be quite easily circumscribed 

and its naturalness has been strongly confirmed by molecular evidence, including large unique deletions 

in two of the sequenced regions (Trift & al. 2004). Recognising Dionysia as a genus, however, 

makes Primula polyphyletic, since the sister group of Dionysia is not Primula, but only a part 

of Primula: the subgenus Sphondylia (Mast & al. 2001, Trift & al. 2004, Wendelbo 1961a-c). 

The species of Primula subg. Sphondylia are similar to some species of Dionysia sect. Dionysiopsis 

Pax (e.g., D. mira and D. bornmuelleri), although the Sphondylia species differ in involute 

young leaves, 3-furrowed pollen grains and a capsule that splits apically into ten teeth. The 

pollen in Dionysia usually has 6-8 furrows. Both pollen types are known from Primula s.str. 

(Wendelbo 1961c). Wendelbo (1961b) considered the subgenus Sphondylia “primitive” in Primula, 

but it seems as if at least the peculiar type of conduplicate-involute leaf vernation (Mast & 

al. 2001) is a derived character (synapomorphy) for the subgenus Sphondylia. 

Towards a natural classification of Dionysia 

It can be concluded from character optimization on phylogenetic trees (Mast & al. 2001, Trift & 

al. 2002, 2004) that the ancestral Dionysia had a lax habit with large dentate revolute leaves, 

flowers in a stalked umbel (or in superposed verticils) subtended by foliaceous bracts and a yellow, 

externally hairy corolla with a long tube – conclusions reached already by Melchior (1943). 

Consequently, there is nothing that suggests that the species with these characteristics (1-5, 

15-18 in the list below) assembled in D. subsect. Scaposae by Wendelbo form a natural group, as 

the traits used to circumscribe subsect. Scaposae are all ancestral in Dionysia. 

Indeed, the molecular results suggest that two “Scaposae” species, Dionysia hissarica and 

D. balsamea, together form a sister group to the other Dionysia. The union of these two species is 

surprising, as it seems to be in conflict with morphological evidence, D. balsamea being superficially 

more similar to D. paradoxa. However, the molecular evidence is very clear and there are 

some morphological characters that distinguish the two from D. paradoxa: comparatively fewseeded 

capsules, blunt teeth on the leaves and long eglandular hairs. 

The western species (15-18) of subsection Scaposae sensu Wendelbo are better put together 

with the similarly western species of subsection Revolutae Wendelbo, with which they share 

many characters. Although here is no molecular support for this grouping, there is nothing that 

contradicts it. The valid name for this taxon is D. sect. Dionysiopsis. 

The second important deviation from previous classifications (Wendelbo 1961, Grey-Wilson 

1989) is the reshuffling of the species of section Dionysia sensu Wendelbo. It is obvious that un-


Table 1. Revised classification of the genus Dionysia. 

A. Eastern species 

I. Probably polyphyletic group not placed to section 

D. hissarica, balsamea, paradoxa, lacei, saponacea 

Large leaves (> 10 mm) with revolute margin 

II. D. sect. Dionysiastrum Smoljan. Flat leaves up to 10 mm 

a. D. subsect. Tapetodes Wendelbo 

D. denticulata, tapetodes [kossinskyi] 

b. D. subsect. Involucratae Wendelbo 

D. involucrata [gandzhinae], hedgei, freitagii, 

viscidula [wendelboi], microphylla 

c. D. subsect. Aghanicae Grey-Wilson 

D. afghanica 

d. D. subsect. Heterotrichae Wendelbo 

D. lindbergii 

Leaf veins raised, farina wolly; corolla yellow, glabrous 

Leaf veins raised, farina powdery (if present); corolla 

violet, glandular-hairy 

Leaves entire, truncate, with obscure veins, covered 

with short glandular hairs; corolla violet, glandularhairy 

Leaves entire, truncate, with obscure veins, covered 

with short glandular hairs and long dense eglandular 

hairs towards the apex; corolla violet, glabrous 

B. Western species 

III. D. sect. Dionysiopsis Pax 

Leaf margin revolute, farina usually present 

D. mira, viva (nov.), bornmuelleri, teucrioides, aretioides, 

leucotricha, revoluta, archibaldii, zschummelii 

(nov.), esfandiarii, rhaptodes, oreodoxa 

IV. D. sect. Zoroasteranthos Lidén (nov.) 

D. curviflora, janthina, khatamii 

V. D. sect. Mucida Lidén (nov.) 

D. lurorum 

VI. D. sect Dionysia 

D. sawyeri (incertae sedis), iransharii, caespitosa, 

gaubae, odora, haussknechtii, crista-galli (nov.), 

zetterlundii (nov.), tacamahaca (nov.), lamingtonii, 

mozaffarianii, iranica, zagrica, khuzistanica, termeana, 

michauxii, bryoides, diapensiifolia, sarvestanica 

Leaves flat, eglandular with long articulate hairs, 

no farina; corolla violet with yellow eye and deeply 

emarginate lobes; seeds few 

Leaves flat, strongly farinose; flowers yellow; seeds 

numerous 

Leaves flat without farina; flowers yellow, or – if 

violet – plant with abundant glandular hairs or corolla 

lobes not emarginate; seeds few 

related elements were brought together here. It is not surprising that the subsections Tapetodes 

and Heterotrichae are shown to be more related to the eastern species of sect. Dionysiastrum as 

they share similarities in leaf shape and leaf anatomy (Bokhari & Wendelbo 1976, Trift & 

Anderberg 2006). Also geographically they come closer. 

More perplexing it is that the densely pulvinate species Dionysia curviflora and D. janthina 

from the Schir Kuh area SW of Yazd (included in D. sect. Dionysia subsect. Bryomorphae by 

Wendelbo) are not related to superficially similar species from the Zagros chain such as D. 

lamingtonii. Instead they group with strong support with the third Schir Kuh endemic, the recently 

described rather lush D. khatamii. If analysed more in detail, this grouping is less surprising; 

all the Schir Kuh species have pubescent violet corollas with a yellow eye and deeply 

emarginate corolla lobes, entire leaves equipped with long coarse articulate hairs and lack glands 

on the leaf lamina. These three species are here recognised as a new section Zoroasteranthos. 

Wendelbo divided the remaining Iranian species with small flat leaves (all from Zagros) between 

the subsections Bryomorphae (i.e. together with the Schir Kuh species) and Caespitosae, 

based on cushion compactness and degree of exsertion of style in pin-eyed flowers. However,


e.g., Dionysia sarvestanica and D. zagrica, agreeing with subsect. Bryomorphae in their compact 

habit, nevertheless have exserted styles, which is a Caespitosae character. It is also clear in the 

molecular phylogeny that the two groups are completely intermingled. 

Three conclusions that can be drawn from the molecular study by Trift & al (2004): (1) geographically 

neighbouring species or higher clades are usually each others closest relatives; (2) 

very tight cushions have evolved four times in dry areas from ancestors with less condensed 

growth; and (3) Dionysia presumably originated in the eastern part of the current distribution 

area. For a fuller discussion of these patterns the reader is referred to that paper. 

Based on the discussion above, I suggest a classification (Table 1), which recognises morphologically 

distinguishable entities that are either well supported in the molecular tree of Trift 

& al (2004), or (in the case of Dionysia sect. Dionysiopsis) is not in conflict with well supported 

nodes in that three. 

Note that the Dionysia subsect. Tapetodes and Heterotrichae have been transferred from D. 

sect. Dionysia to sect. Dionysiastrum. 

Key to the species of Dionysia 

1. Leaf margin revolute or all leaves more than 10 mm. . . . . . . . . . . . . . . . 2 

– Leaves flat or slightly involute, 2-10 mm (bracts and leaves of new “vegetative” shoots occasionally 

longer) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18 

2. Flowers purple, violet or pink . . . . . . . . . . . . . . . . . . . . . . . . 3 

– Flowers yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5 

3. Leaf indumentum very dense, apically retrorse; corolla hairy; cushions very dense . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26. D. esfandiarii 

– Leaf indumentum sparse to moderately dense, not retrorse apically; corolla glabrous or almost 

so . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4 

4. Cushions very dense; hairs few and short; leaf margin entire, strongly inrolled . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23. D. zschummelii 

– Cushions lush; leaves distinctly hairy, crenate-dentate . . . . . . . . 22. D. archibaldii 

5. Inflorescence consisting of an umbel or of superposed verticils of flowers . . . . . . 6 

– Flowers sessile, 1-2 per inflorescence; bracts linear . . . . . . . . . . . . . . . 13 

6. Inflorescence sessile . . . . . . . . . . . . . . . . . . . . . . . . 4. D. lacei 

– Inflorescence stalked, bracts foliaceous. . . . . . . . . . . . . . . . . . . . . 7 

7. Leaves 25-80 mm, inflorescence usually with 2 or more whorls of flowers, one above the 

other . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8 

– Leaves 10-20 mm, inflorescence a single umbel with 2-3 flowers . . . . . . . . . 12 

8. Leaves with short-stalked glands only . . . . . . . . . . . . . . . 3. D. paradoxa 

– Leaves with both long hairs and short-stalked glands . . . . . . . . . . . . . . . 9 

9. Capsule with 5-10 seeds; corolla limb 10-18 mm broad, with broad obovate lobes (Afghanistan) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. D. balsamea 

– Capsule with 40-70 seeds; corolla limb 7-10 mm broad, lobes broadest towards the base, 

rectangular to oblong-triangular . . . . . . . . . . . . . . . . . . . . . . . 10 

10. Leaves 5 times as long as broad, with 10-15 regular teeth on each side . . . . 15. D. mira 

– Leaves 3 times as long as broad, with 4-5 irregular teeth or lobes . . . . . . . . . 11 

11 Corolla tube 20-30 mm; calyx split to 3 /4 or more . . . . . . . . . 16. D. bornmuelleri 

– Corolla tube 13-15 mm; calyx split to 2 /3 . . . . . . . . . . . . . . . 17. D. viva 

12. Leaves clearly revolute . . . . . . . . . . . . . . . . . . . . 18. D. teucrioides 

– Leaves almost flat . . . . . . . . . . . . . . . . . . . . . . . 1. D. hissarica 

13. Leaves 15-30 × 6-20 mm, flat at maturity . . . . . . . . . . . . . 5. D. saponacea 

– Leaves much shorter and narrower, strongly revolute . . . . . . . . . . . . . . 14 

14. Corolla lobes emarginate . . . . . . . . . . . . . . . . . . . . . . . . . . 15 

– Corolla lobes entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17


15. Calyx lobes with broad blunt erose-dentate apices . . . . . . . . . 20. D. leucotricha 

– Calyx lobes entire, acute . . . . . . . . . . . . . . . . . . . . . . . . . . 16 

16. Leaves densely glandular with 5-8 crenations . . . . . . . . . . . . 21. D. revoluta 

– Leaves sparsely glandular with 4-5 crenations . . . . . . . . . . . 19. D. aretioides 

17. Cushions very dense; leaves bluish green with densely set short, acute hairs above (rarely 

glabrous) and stalked glands marginally and below; leaf margin usually strongly inrolled, 

not or only very slightly crenate-dentate . . . . . . . . . . . . . 24. D. rhaptodes 

– Cushions soft; leaves bright green, glabrous or with a few scattered hairs, with subsessile 

glands marginally and below, usually farinose; leaf margin clearly crenate-dentate . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25. D. oreodoxa 

18. Flowers in a stalked umbel (rarely with a single flower); bracts foliaceous, usually larger 

than leaves . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19 

– Flowers sessile, single or rarely in pairs; bracts shorter and much narrower than leaves 24 


– Flowers yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22 

20. Leaves 1.5-2.5 mm, with few veins . . . . . . . . . . . . . . . 13. D. microphylla 

– Leaves 5-12 mm, with conspicuous flabellate venation . . . . . . . . . . . . . . 21 

21. Leaves and bracts dentate; corolla lobes emarginate; homostylous . . 12. D. involucrata 

– Leaves and bracts entire; corolla lobes acute, entire; heterostylous . . . . 11. D. hedgei 

22. Plant not farinose or only on corolla; leaves in a tight rosette . . . . 33. D. caespitosa 

– Plant farinose (especially in axils); leaves not in a tight rosette . . . . . . . . . . 23 

23. Corolla hairy; leaves crenate-lobed . . . . . . . . . . . . . . . . 1. D. hissarica 

– Corolla glabrous; leaves entire to serrate . . . . . . . . . . . . . . 30. D. lurorum 

24. Strongly anisophyllous with “vegetative” leaves 3-4 times longer than overwintering 

leaves; corolla glabrous, lobes deeply emarginate; ovules 15-25 (corolla limb possibly violet/purple) 

. . . . . . . . . . . . . . . . . . . . . . . . . . 31. D. sawyeri 

– Not strongly anisophyllous; corolla usually hairy; ovules 3-10 . . . . . . . . . . 25 


– Flowers yellow . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 35 

26. Leaves with strongly raised veins . . . . . . . . . . . . . . . . . . . . . . 27 

– Leaf veins not conspicuous . . . . . . . . . . . . . . . . . . . . . . . . . 28 

27. Calyx 5 mm, divided to 1 /2; corolla tube c. 15 mm . . . . . . . . . . 10. D. freitagii 

– Calyx 3-4 mm, divided to 3 /4; corolla tube 10 mm . . . . . . . . . . 9. D. viscidula 

28. Leaves apically divided into 3-5 lobes . . . . . . . . . . . . . . 41. mozaffarianii 

– Leaves entire . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29 

29. Leaves with long eglandular hairs (0.5 mm), with or without short glandular hairs . . . 30 

– Leaves without eglandular hairs but densely set with sessile or short-stalked glands . . 34 

30. Leaves with short-stalked glands; calyx split to base . . . . . . . . 14. D. lindbergii 

– Leaves without glands; calyx divided 1 /2 - 2 /3 . . . . . . . . . . . . . . . . . . 31 

31. Leaves obovate, glabrous below . . . . . . . . . . . . . . . . . 27. D. curviflora 

– Most leaves hairy on both surfaces . . . . . . . . . . . . . . . . . . . . . . 32 

32. Corolla glabrous . . . . . . . . . . . . . . . . . . . . . . . 32. D. iranshahrii 

– Corolla hairy . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33 

33. Leaves c. 5-8 mm, cushions loose . . . . . . . . . . . . . . . . . 29. D. khatamii 

– Leaves 3 mm, cushions dense . . . . . . . . . . . . . . . . . . 28. D. janthina 

34. Calyx split to base, leaves narrowly elliptic . . . . . . . . . . . . . 47. D. bryoides 

– Calyx split to 2 /3, leaves rounded-obtriangular . . . . . . . . . . . 6. D. afghanica 

35. Leaves without glands or glandular hairs . . . . . . . . . . . . . . . . . . . 36 

– Leaves glandular hairy or with sessile glands . . . . . . . . . . . . . . . . . . 37 

36. Leaves with long (0.5 mm) curly articulate hairs; corolla glandular hairy . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 40. D. lamingtonii


– Leaves with short straight retrorse hairs; corolla with eglandular hairs . 46. D. michauxii 

37. Corolla glabrous, very rarely with short eglandular hairs (NE Iran to Afghanistan) . . 38 

– Corolla glandular hairy (Zagros mountains) . . . . . . . . . . . . . . . . . . 39 

38. Calyx lobes mucronate . . . . . . . . . . . . . . . . . . . . . 7. D. denticulata 

– Calyx lobes not mucronate . . . . . . . . . . . . . . . . . . . . 8. D. tapetodes 

39. Leaves of adult plants up to 3 mm, entire or rarely slightly tridentate at apex . . . . 40 

– Leaves longer, at least some with a few lateral teeth or lobes . . . . . . . . . . . 46 

40. Leaves with a uniform indumentum of sessile or short-stalked glandular hairs (up to 0.1 mm) 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 41 

– Leaves with some longer hairs, especially adaxially . . . . . . . . . . . . . . . 43 

41. Glands sessile (leaves obovate, calyx lobes broadest at the middle) . . . 43. D. zagrica 

– Glands shortly stalked . . . . . . . . . . . . . . . . . . . . . . . . . . . 42 

42. Leaves oblong to obovate, rounded obtuse at apex; calyx lobes usually broadest at the middle 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . 49. D. sarvestanica 

– Leaves ovate, often tridentate at apex; calyx lobes broadest at the base . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 44. D. khuzistanica 

43. Calyx divided to about halfways; leaves acute, densely glandular hairy on both sides . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39. D. tacamahaca 

– Calyx divided almost to the base; leaves obtuse . . . . . . . . . . . . . . . . 44 

44. Leaves with an apical whitish fimbriate crest . . . . . . . . . . . 37. D. crista-galli 

– Apical marginal hairs not confluent to a whitish crest . . . . . . . . . . . . . . 45 

45. Leaves with ciliate margin; most hairs gland-tipped . . . . . . . 36. D. haussknechtii 

– Leaf margin not distinctly ciliate; leaves adaxially with long straight eglandular hairs in addition 

to shorter glandular hairs . . . . . . . . . . . . . . . . 38. D. zetterlundii 

46. Leaves with eglandular hairs more than 0.5 mm . . . . . . . . . . . . . . . . 47 

– All hairs gland-tipped, less than 0.2 mm . . . . . . . . . . . . . . . . . . . 48 

47. Calyx 5-6 mm; marcescent leaves not spirally reflexed . . . . . . . . . 35. D. odora 

– Calyx 7.5-9 mm; marcescent leaves spirally reflexed . . . 34. D. gaubae v. megalantha 

48. Glands all sessile . . . . . . . . . . . . . . . . . . . . . . . . 42. D. iranica 

– Most glands clearly stalked . . . . . . . . . . . . . . . . . . . . . . . . . 49 

49. Calyx divided to 3 /4; marcescent leaves strongly reflexed so as to become more or less 

coiled . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34. D. gaubae 

– Calyx divided almost to the base; marcescent leaves not reflexed . . . . . . . . . 50 

50. Style of long-styled flowers long exserted; calyx accrescent to 10 mm in fruit . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 48. D. diapensiifolia 

– Style in long-styled flowers not or only slightly exserted; calyx c. 5 mm in fruit . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45. D. termeana 

List of taxa 

Synonyms are listed in chronological order, except that homotypic synonyms are kept together. 

The list is (hopefully) nomenclaturally complete, although the emphasis is on Iranian species. 

Descriptions are restricted to new taxa plus a few corrections and emendations to the previous 

treatments by Wendelbo (1961), Grey-Wilson (1989) and Jamzad (1996). Three eastern species, 

for which taxonomic status has not been ascertained as the types have not been seen, are included 

in the list under a closely similar species. 

Dionysia Fenzl in Flora 26: 389. 1843 ≡ Primula sect. Dionysia (Fenzl) Kuntze in Post & Kuntze, 

Lex. Gen. Phan.: 406. 1903. – Type: D. odora Fenzl 

= Macrosyphonia Duby in Mem. Soc. Phys. Hist. Nat. 10: 426. 1844. – Type: Dionysia caespitosa 

(Duby) Boiss. [≡ Gregoria caespitosa Duby ≡ Macrosyphonia caespitosa (Duby) Duby].


A. Eastern species 

I. Species not placed to section 

1. Dionysia hissarica Lipsky in Trudy Imp. St.-Petersburgsk. Bot. Sada 18: 83. 1900 ≡ Primula 

hissarica (Lipsky) Bornm. in Bull. Herb. Boissier, ser. 2, 3: 592. 1903. – Holotype: USSR, 

Pamir-Alai, above Den-Surkh, near river Surkhan, Lipsky (LE; isotypes: A, B, E, G) 

Morphologically, the most aberrant species in the genus. A possibly unique feature not noted 

before is the finely papillose spherical seeds. Dionysia balsamea (its putative sister species, Trift 

& al. 2004) has, however, not been studied in this respect. If they share this character, it would constitute 

a strong corroboration of the molecular tree. 

2. Dionysia balsamea Wendelbo & Rech. f. in Aarbok Univ. Bergen Mat.-Naturvitensk. Ser. 

19(4): 7, fig. 2, 3e-f. 1964. – Holotype: Afghanistan, Ghorat, Kuh-Tscheling-Safed-Daraq (Pirestan), 

7.-8.1962, Rechinger 19094 (W;isotype:BG). 

3. Dionysia paradoxa Wendelbo in Bot. Not. 112: 497. 1959. – Holotype: Afghanistan, Kabul 

river, Sarobi, Volk 2411 (BG; isotype: BASBG). 

4. Dionysia lacei (Hemsley & Watt) Clay, Present-Day Rock Gard.: 195. 1937 ≡ Primula lacei 

Hemsley & Watt in J. Linn. Soc. Bot. 28: 298, 325, t. 41. 1891. – Holotype: Pakistan, Baluchistan, 

Torkhan, Lace 3648 (K; isotypes: BM, E). 

5. Dionysia saponacea Wendelbo & Rech. f. in Aarbok Univ. Bergen Mat.-Naturvitensk. Ser. 

19(4): 10, fig. 3a-b. 1964. – Holotype: Afghanistan, Ghorat, Kuch-Tscheling-Safed-Daraq (Pirestan), 

7.-8.1962, Rechinger 19092 (W; isotype BG). 

II. Dionysia sect. Dionysiastrum Smoljan. in Siskin & Bobrov, Fl. SSSR 18: 718. 1952. – Type: 

D. involucrata Zaprjag. 

a. Dionysia subsect. Afghanicae Grey-Wilson, Gen. Dionysia: 168. 1989. – Type: D. afghanica 

Grey-Wilson 

6. Dionysia afghanica Grey-Wilson in Kew Bull. 29: 57. 1974. – Holotype: Afghanistan, Darreh 

Zang, S of Belcheragh, 7.1971, Grey-Wilson & Hewer 1308 (K; isotypes: E, GB, W). 

b. Dionysia subsect. Tapetodes Wendelbo in Aarbok Univ. Bergen Mat.-Naturvitensk. Ser. 

19(4): 10. 1964 [– D. sect. Bryonanthe Smoljan., nom. nud.]. – Type: Dionysia tapetodes Bunge 

7. Dionysia denticulata Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 19(4): 11, fig. 4. 

1964. – Holotype: Afghanistan, Bamian, between Shahtu and Panjao, 6.1962, Hedge & Wendelbo 

W 4876 (BG; isotype: E). 

8. Dionysia tapetodes Bunge in Bull. Acad. Imp. Sci. Saint-Pétersbourg 16: 562. 1871 ≡ Primula 

tapetodes (Bunge) Kuntze, Rev. Gen. Pl. 2: 400. 1891. – Holotype: NE Iran, above Derrud, between 

Nishapur and Meshed (Masshad), Bunge (LE; isotypes: G, P). 

= Dionysia trinervia Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 19(4): 14. 1964. – Holotype: 

Afghanistan, Orozgan, Kotal-Tachakmak, 3100 m, 6.1960, K. Lindberg 938 (BG). 

8b. Dionysia kossinskyi Czerniak. in Izv. Glavn. Bot. Sada SSSR 26: 116. 1927. – Holotype: 

Iran: Khorrasan, Mt. Kisil-Chisht, above Khorkei, Czerniakovska 375 (LE). 

The type (and Czerniakovska 96, LE, from the same area) differs from “typical” Dionysia 

tapetodes in having a corolla with short eglandular hairs, and leaves with slightly longer glands. 

Possibly, it can also be recognised by longer leaves and pink or violet corolla, but this is as yet uncertain.


c. Dionysia subsect. Involucratae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 78. 

1961. – Type: D. involucrata Zaprjag. 

= Dionysia subsect. Microphyllae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 78. 

1961. – Type: D. microphylla Wendelbo 

9. Dionysia viscidula Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 19: 20, fig. 11, 16. 1964. 

– Holotype: Afghanistan, Darrah Zang, near Belcheragh, 5.1962, Hedge & Wendelbo W 3722 (BG; 

isotype: E). 

9b. Dionysia wendelboi Podlech in Mitt. Bot. Staatssamml. München 17: 481. 1981. 

“Differt a Dionysia viscidula foliis minoribus farinosis minutissime tantum glandulosipuberulis 

floribus minoribus”. I have not seen the type of this name. The species is said to differ 

from D. viscidula in smaller leaves and flowers. 

10. Dionysia freitagii Wendelbo in Bot. Not. 123: 303, fig. 2A-F. 1970. – Holotype: Afghanistan, 

Balkh, Ali Kuh, 5.1969, Hedge, Wendelbo & Ekberg W 8497 (GB; isotype: E). 

11. Dionysia hedgei Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 19: 18, fig. 9, 10-16. 

1964. – Holotype: Afghanistan, Mazar-i-Sharif, Koh-i-Elburz, Hedge & Wendelbo W3888 (BG; 

isotype: E). 

12. Dionysia involucrata Zaprjag. in Trudy Tadzhiksk Bazy, Bot. 2: 153, fig. 3. 1936. – Holotype: 

USSR, Pamir-Alai, Hissar (LE). 

12b. Dionysia gandzhinae Kamelin in Bot. Zhurn. 69(10): 1403. 1984. – Holotype: S Tadjikistan, 

jugum Gasimailik, declivum orientale, prope pagum Gandzhina et supra locum Gusabulak, 

in fissuris rupium 3.5.1983, R. Kamelin, A. Borodina, A. Geldychanov, I. Mochova, E. Nikolaev, 

T. Schevljakova, V. Solovjov & I. Tagaev N 1437 (LE?, TAD, not seen). 

Although the type of Dionysia gandhzinae has not been seen (it could not be recovered in LE), 

it must, judging from the description, be very similar to D. involucrata. Alleged points of difference 

are the shorter peduncles (“1 cm” instead of 1.2-1.8) and smaller flowers (“15-18 × 7-8 mm” 

instead of 20-29 × 9-14 mm). 

13. Dionysia microphylla Wendelbo in [Köie & Rech. f., Symb. Afghan. 4] Biol. Skr. 10(3): 68, 

fig. 26. 1958. – Holotype: Afghanistan, around 35°N, 65°E, Edelberg 2313 (C;isotype:BG). 

Contrary to the descriptions by Wendelbo and Grey-Wilson, the leaves usually have short 

hairs towards the base, not only sessile glands. 

d. Dionysia subsect. Heterotrichae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 78. 

1961. – Type: D. lindbergii Wendelbo 

14. Dionysia lindbergii Wendelbo in Bot. Not. 112: 495, fig. 1. 1959. – Holotype: Afghanistan, 

Darreh Zang, Lindberg 454 (BG; isotype: LD). 

B. Western species 

III. Dionysia sect. Dionysiopsis (Pax) Melchior in Mitt. Thüring. Bot. Vereins 50: 159. 1943 ≡ 

Primula sect. Dionysiopsis Pax in Jahresber. Schles. Ges. Vaterl. Cult. 87: 20. 1909 ≡ Dionysia 

sect. Ariadna Wendelbo in Bot. Not. 112: 496. 1959, nomen superfl. ≡ Dionysia sect. Anacamptophyllum 

subsect. Scaposae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 77. 1961. – 

Type: D. bornmuelleri Pax 

= Dionysia sect. Anacamptophyllum Melchior in Mitt. Thüring. Bot. Vereins 50: 167. 1943. – 

Type: D. aretioides Lehm. 

= Dionysia sect. Anacamptophyllum subsect. Mirae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. 

Ser. 3: 77. 1961. – Type: D. mira Wendelbo 

= Dionysia sect. Anacamptophyllum subsect. Revolutae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. 

Ser. 3: 77. 1961. – Type: D. revoluta Boiss.


15. Dionysia mira Wendelbo in Bot. Not. 112: 500. 1959 ≡ Primula aucheri Jaub. & Spach, Ill. 

Pl. Orient. 1: 97, t. 49. 1842. – Holotype: Oman, Djebel Akdar, Aucher-Eloy 5236 (P; isotypes: 

BM,G,K,UPS)–nonDionysia aucheri (Duby) Boiss., q.e. D. odora. 

Grey-Wilson (1989) gives the calyx division as halfways. However, usually it is divided to at 

least 3 /4. Although this species is similar in detail to Dionysia bornmuelleri, it has a rather different 

habit. 

16. Dionysia bornmuelleri (Pax) Clay, Present-Day Rock Gard.: 194. 1937 ≡ Primula bornmuelleri 

Pax in Jahresber. Schles. Ges. Vaterl. Cult. 87: 21. 1909. – Holotype: Iran, Kermanshah, 

Noah Kuh near Kerind, Strauss 601 (B; isotypes: E, K, W). 

In Grey-Wilson (1989) this species is said to have a smaller petal limb than Dionysia mira.At 

least as often the opposite relation holds. 

17. Dionysia viva Lidén & Zetterlund, sp. nov. 

Holotype: Iran, Fars, Vallyabad (29.47.19; 53.12.56), 1700 m, 22.4.2002, Zschummel, Zschummel, 

Zetterlund & Tjerdsmaa T4Z 035 (GB; isotypes: TARI, UPS) – Fig. 1. 

Caespites laxi. Folia (petiolo incluso) 25-50 × 7-13 mm, lamina obovata apice rotundata, in 

petiolum anguste alatum attenuata, margine plus minusve profunde sinuato-dentata vel duplodentata, 

subtus lanato-farinosa, pilis glandulosis et eglandulosis obsita. Inflorescentia everticillis 

superpositis 1-2(-3)-nis approximatis 2-4-floris, scapo (2-)5-8 cm longo. Bracteae ovatae 

12-18 × 4-9 mm, grosse serratae. Calyx 9-10 mm longus, ad 2 /3 in segmenta late lanceolata divisus. 

Corolla (longistylis) 14 mm longa, flava, pilis crispatis glandulosis obsita, limbo 5-6 mm 

diametro lobis 2-2.5 mm longis subrectangularibus. Capsula c. 70-sperma, seminibus atrobrunneis 

0.5 mm longis subquadrangularibus. 

Fig. 1. Dionysia viva – A: calyx; B: corolla, showing size extremes in cultivation; C: whole plant; D-F: hair 

details, corolla (D), calyx (E), underside of leaf (F). – From the holotype (C) and flower details (A, D-F)from 

plants cultivated in the Göteborg Botanic Garden from seeds of the holotype.


Dionysia bornmuelleri Pax calyce fere ad basim diviso lobis angustioribus, tubo corollae profunde 

longiore, a D. viva differt. 

Etymology. – Swedish “viva” means any species of the genus Primula. 

Lax chasmophytic subshrub with leaves crowded at branch tips; each terminal rosette with 1-2 

sparsely leafy “vegetative” axillary shoots, 2-6 cm long, eventually producing new terminal rosettes 

towards the end of the season, and with one flowering scape (rarely with second scape); 

defoliated stem segments with a reddish brown lustre; the leaf (remnants) crowded towards the 

end of the branches (stem segments) clearly discernible also in older parts; the amount of secondary 

growth modest, thickest stems 3-4 mm in diameter. Indumentum of green plant parts of 

(a) scattered sessile glands, (b) rather dense many-celled articulated glandular hairs 0.2-1 mm 

long, and especially on veins on the underside of leaves, (c) scattered coarse and straight, acute 

eglandular hairs up to 1.5 mm. Farina woolly and mainly on the underside of leaves, at petiole 

bases and inside of calyx. Leaves (including petiole) 25-50 × 7-13 mm, narrowed into a long narrowly 

winged petiole, lamina obovate, with raised veins below, flat or (especially when young) 

with slightly revolute margin, coarsely dentate with obtuse to subacute primary teeth, each usually 

with 1-3 minute secondary teeth. Scapes (2-)5-8 cm long with 1-3 closely set whorls of flowers, 

each with 2-4 flowers. Bracts ovate, sessile, coarsely serrate, 9-17 × 4-9 mm (or the uppermost 

smaller). Calyx 9-10 mm, divided to 2 /3 into broadly oblanceolate lobes. Corolla (longstyled) 

yellow, sparsely crispate-glandular hairy in lower half, more densely so in upper half and 

on the underside of lobes; tube 14 mm long, limb 5-6 mm broad with lobes 2 mm long, obtusely 

sub-rectangular in shape, sometimes with a minute notch at apex; style in long-styled flowers 

exserted. Capsule rounded with up to 70 dark brown small angular seeds, 0.5 mm long. 

Dionysia viva is vegetatively very similar to D. bornmuelleri, only somewhat less hairy and a bit 

smaller in all parts. It is, however, well separated on floral characters. In D. bornmuelleri the calyx 

is split almost to the base and the calyx lobes are narrow, the corolla tube is much longer 

(20-30 mm; in D. viva 14 mm) and the corolla lobes are differently shaped, being bluntly triangular. 

D. bornmuelleri is distributed far to the NW of D. viva. 

The plants of Dionysia viva in cultivation are, as can be expected, larger in most vegetative 

parts, and the floral whorls are more distant. They show a surprising variation in floral width; one 

plant had flowers twice as broad as those of the type. 

18. Dionysia teucrioides P. H. Davis & Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 76. 

1961. – Holotype: Turkey, Hakkari, Cilo Dag, in Diz derezi, 6.8.1954, P. H. Davis & Polunin 

23884 (K;isotype:E). 

19. Dionysia aretioides (Lehm.) Boiss., Diagn. Pl. Orient., ser. 1, 7: 68. 1846 ≡ Primula aretioides 

Lehm., Monogr. Primul.: 90, t. 9. 1817 ≡ Gregoria aretioides (Lehm.) Duby in Candolle, Prodr. 8: 

46. 1844. – Holotype: N Iran, Ghilan, Hablitz (LE?;isotypes:B,BN,C,G,W). 

= Dionysia demawendica Bornm. in Beih. Bot. Centralbl., Abt. 2, 33: 301, t. 2 fig. 1. 1915. – 

Holotype: N Iran, foot of Mt Demawand, Abigerm, Bruns (B; not seen, teste Wendelbo). 

20. Dionysia leucotricha Bornm. in Beih. Bot. Centralbl., Abt. 2, 28: 460. 1911 ≡ Dionysia aretioides 

var. adenophora Bornm. in Bull. Herb. Boissier, ser. 2, 3: 593, t. 6 fig. 5. 1903. – Holotype: 

Iran, Mt Elvend by Hamadan, Strauss (B;isotype:JE). 

Superficially similar to Dionysia aretioides and D. revoluta, but immediately distinguished by 

the peculiar calyx with long hairs at base and broad dentate lobe-apices. 

21. Dionysia revoluta Boiss., Diagn. Pl. Orient., ser. 1, 7: 65. 1846 ≡ Primula revoluta (Boiss.) 

Bornm. in Bull. Herb. Boissier 7: 73. 1899. – Holotype: Iran, Fars, Kuh-e-Sabzpuchon, Kotschy 

426 (G; isotypes: B, BM, K, W). 

= Dionysia revoluta var. canescens Boiss., Fl. Orient. 4: 18. 1879 ≡ Dionysia revoluta subsp. 

canescens (Boiss.) Wendelbo in Aarbok Univ. Bergen Mat.-Naturvitensk. Ser. 3: 48. 1961. –


Lectotype (designated by Wendelbo, id.): Iran, Luristan, Kuh Eschker, Haussknecht (G [not 

seen]). 

This is the most common and widespread of the Dionysia species in Zagros. Although it is variable, 

it is questionable whether subsp. canescens can be upheld despite geographical trends. In the 

type locality, however, there are plants with dense long hairs occurring together with bright green 

individuals with only glandular hairs. Long eglandular hairs are frequently found also on the corolla. 

22. Dionysia archibaldii Wendelbo in Bot. Not. 120: 144. 1967. – Holotype: Iran, Bakhtiari, 

Tang-i-Sirdan, 4000 m, 8.1966, Archibald 3053 (GB). 

= Dionysia bazoftica Jamzad in Iran. J. Bot. 7: 20. 1996. – Holotype: Bakhtiari, Bazoft, Chebd, N 

slope of Kuh-e-Taraz, 1700-2300 m, Mozaffarian 57824 (TARI). 

The seedlings raised from the original Archibald collection gave rise to a very heterogeneous 

offspring (Grey-Wilson 1989: 90), as is indeed very common with Dionysia. The type material 

(GB, collected in the field) has broad, crenate-dentate leaves with long hairs and rather lush 

growth. The single Archibald clone still surviving in cultivation, however, is of a denser type 

with less hairy and strongly revolute almost entire leaves. It should be noted that at least part of 

Archibald’s original introduction was collected at very high altitude late in the year. 

After having seen a good number of individuals in the field, in cultivation, in herbaria and on 

photographs I have come to the conclusion that this and the following taxon are clearly separate 

species. In leaf morphology, they afford a striking parallel to the species pair D. rhaptodes/oreodoxa 

(24-25). Unlike these two they seem to have non-overlapping, though contiguous, distribution 

areas. 

Jamzad (1996) described a new species, Dionysia bazoftica, said to differ from D. archibaldii 

i.a. in the possession of long hairs (she also lists differences in leaf arrangement, and a few 

other minor details). However, the type specimen of D. archibaldii indeed has long hairs, lush 

growth, rather broad, strongly crenate leaves, etc., and is in every respect similar to the type of D. 

bazoftica, which must accordingly be sunk in synonymy. 

A population from the upper Kuhrang river was studied more closely by myself in 1998. The 

plants here vary a lot in indumentum: most have long hairs, but there are also some without. Presence 

of farina also varies within populations. General habit is partly depending on exposure, age 

and time of the year, but there is a very strong genetic component. In seed offspring from a single 

plant of this population (SLIZE 115, cult. Göteborg Botanic Garden) one is struck by the immense 

variation shown (much surpassing that of adult plants in the wild), both in vegetative and floral 

characters, clearly showing the high degree of heterozygosity in individuals of these populations, 

as indeed in many other species, such as Dionysia aretioides, D. termeana and D. diapensiifolia. 

Considerable variation in populations of D. archibaldii was noted by the T4Z expedition. 

According to Grey-Wilson (1989) the corolla is sometimes slightly glandular-pubescent. The 

number of ovules are 4-8, but often only 3-4 seeds develop. 

23. Dionysia zschummelii Lidén, sp. nov. 

Holotype: Iran, Lorestan, between Aligoudarz and Shoulabad, Ghadee Kuh (33.06.42; 49.25.25), 

2600-2700 m, 5.5.2002, T4Z 166 (UPS; isotypes: GB, TARI) – Fig. 2. 

A Dionysia archibaldii habitu densissimo, foliis minutis crassiusculis fere glabris differt. 

Caespites densissimi non farinosi. Folia parva, sessilia, lamina oblonga obtusa margine non 

crenato-dentata, profunde et abrupte revoluta, fere glabra. Bracteae lineares. Calyx 3.5-4 mm 

longus, usque ad basin in lobis linearibus divisus. Corolla 10-14 mm longa, purpureo-lilacina vel 

lavandulacea, glabra, limbo 9-14 mm diametro lobis late ovatis vel obovatis profunde emarginatis 

stylo parce excerto. Capsula c. 4-sperma, seminibus atrobrunneis 0.9 mm longis. 

Eponymy. – Named after Dieter Zschummel, who discovered this species and brought it into cultivation.


Fig 2. A-D: Dionysia zschummelii – corolla (A); calyx with bract (B); leaf, abaxial view (C); flowering 

branch (D); from plants cultivated in the Göteborg Botanic Garden from seeds of the type collection. – E-G: D. 

archibaldii – corolla (E); calyx segment (F); flowering branch (G); upper Kuhrang valley, SLIZE 115, UPS. 

Very dense non-farinose cushions. Leaves 5-6 × 1.5 mm, bluish green, sparsely and shortly 

hairy; slightly longer hairs on the under side, but due to the sharply reflexed leaf margin these 

not visible. Bracts 2, 3.5-4 mm long, linear, shortly hairy abaxially, glabrous adaxially. Calyx 

4(-4.5) mm, divided to the base into linear obtuse erect lobes, shortly hairy outside, glabrous inside. 

Corolla glabrous, pale lilac to lavender; tube 10-11 mm long; limb 10-12 mm across, divided 

into ovate rather deeply emarginate lobes. Style of long-styled flowers slightly exserted. 

Dionysia zschummelii is a northern vicariant of D. archibaldii with extremely dense habit. It has 

small entire leaves with strongly inrolled leaf margins, so as to render the leaf subtrigonous-semicircular 

in cross-section. The few hairs are mainly confined to the edges where the pale greyish 

green thickish central plain of the upper leaf surface is more or less sharply reflexed into a 

darker more bluish green lateral side. The flowers are very similar to those of D. archibaldii. 

Plants in cultivation differ but little from those of wild populations. 

Additional record. – Type locality: 7.5.2001, Dionysia zschummel 01-17 (UPS). 

24. Dionysia rhaptodes Bunge in Bull. Acad. Imp. Sci. Saint-Pétersbourg 16: 562. 1871. – Holotype: 

Iran, Kerman, between Chabbis and Kerman, 30.3.1859, Bunge (E; isotypes: G, K, P). 

= Dionysia heterochroa Bornm. in Bull. Herb. Boissier 7: 72, t. 2 fig. 3. 1899 ≡ Primula heterochroa 

(Bornm.) Bornm. in Bull. Herb. Boissier 7: 73. 1899. – Holotype: Kuh-i-Jupar, Bornmüller 

3872 (B; isotypes: G, JE). 

25. Dionysia oreodoxa Bornm. [Primula oreodoxa (non Franchet) Bornm. in nota] in Bull. Herb. 

Boissier 7: 68, t. 2 fig. 1. 1899 ≡ Primula kermanensis Bornm.inBull.Herb.Boissier,ser.2,3: 

592. 1903 – Holotype: Iran, Kerman, Kuh-i-Jupar, Bornmüller 3873b (B). 

26. Dionysia esfandiarii Wendelbo in Bot. Not. 123: 302, fig. 2m-n. 1970. – Holotype: Iran, 

Abadeh, Bacanat, Kuh Khataban, 6.1969, Termé 8128E (GB; isotype: TARI). 

This species is well described by Grey-Wilson (1989) although he complained that the material 

was immature. Material from the SLIZE expedition is now in cultivation in several places. It


lacks glandular hairs and is covered in long articulate hairs that are retrorse towards the apex of 

the leaf. The reflexed leaf margin is often slightly crenate-dentate, indicating relation with the 

other species in the section despite its widely divergent habitus, like a tight hairy ball. 

IV. Dionysia sect. Zoroasteranthos Lidén, sect. nov. 

Type: Dionysia curviflora Bunge 

Plantae plus minusve caespitosae. Folia parva integra pilis articulatis longis non glanduliferis. 

Bracteae parvae. Flores singulares sessiles. Corolla lilacina plerumque flavoannulata lobis emarginatis 

stylo incluso. Semina pauca sat magna. 

Etymology. – Zoroaster, Farsi Zarathushtra, the founder of mazdaism, a very old belief system 

which in Iran has (had) a strong foothold in the Yazd area; Greek: anthos =flower. 

27. Dionysia curviflora Bunge in Bull. Acad. Imp. Sci. Saint-Pétersbourg 16: 562. 1871 ≡ 

Primula curviflora (Bunge) Kuntze, Rev. Gen. Pl. 2: 400. 1891. – Holotype: Iran, Yazd, Schir 

Kuh, Buhse 1352 (G; isotypes: B, P). 

The corolla tube usually has a mixture of tiny eglandular and glandular hairs, rarely glabrous. 

28. Dionysia janthina Bornm. & Winkl. in Bull. Herb. Boissier 7: 70, t. 2 fig. 2. 1899 ≡ Primula 

janthina (Bornm. & Winkl.) Bornm. in Bull. Herb. Boissier 7: 73. 1899. – Holotype: Iran, Yazd, 

[S of] Schir Kuh [proper], Bornmüller 3869 (B; isotypes: BM, E, JE, K). 

29. Dionysia khatamii Mozaff. in Pakistan J. Bot. 34: 391. 2002. – Holotype: Yazd, Mehriz, 

Darre Damgahan, in deep granitic stone wall, 2400-2550 m, V. Mozafarrian 79250 (TARI; 

isotype: UPS). 

This species was recently found and described by Valiollah Mozaffarian, and is in cultivation 

in a few gardens. It is similar to Dionysia iranshahrii in its narrow, densely pubescent leaves and 

violet flowers. It differs, however, in its less dense growth, much larger flowers with deeply divided 

calyx and narrower deeply emarginate petal lobes, and the presence of glands on the stem. 

It is not restricted to granitic rocks, but occurs on limestone as well. 

V. Dionysia sect. Mucida Lidén, sect. nov. 

Type: Dionysia lurorum Wendelbo 

Herba perennis non pulvinaris dense farinosa. Folia late oblanceolata serrata vel integra. Inflorescentia 

scaposa 2-4-flori. Bracteae grandes, saepe profunde serrato-dentatae. Flores lutei. Semina 

minuta numerosa (ad 100!). 

Etymology. – From Latin mucidus = mouldy, referring to the woolly farina that usually covers 

much of the plant. 

30. Dionysia lurorum Wendelbo in Notes Roy. Bot. Gard. Edinburgh 38: 105. 1980. – Holotype: 

Iran, Luristan, 61 km on road from Aligoudarz to Shoulabad (valley after the pass), 2400 m, 

29.6.1977, Runemark & Lazari 26216 (G; isotypes: E, TARI). 

= Dionysia aubrietioides Z. Jamzad & Mozaff. in Iran. J. Bot. 7: 19. 1996. – Holotype: Bakhtiari, 

Bazoft valley, Mavaraz, Talkhdan, Kuh-e-Mafaron (Kuh-e-Sefid) 2700 m, Mozaffarian 74002 

(TARI). 

A homogenous and easily recognised species, very different from all other Dionysia. D. 

aubretioides was based on a shade-grown rather luxurious individual, but is in all details indistinguishable, 

and cultivated plants from the type locality do not differ from those from type locality 

of D. lurorum. 

This species is sometimes completely covered in farina. Shade-grown plants are notable for 

their long and thin internodes, whereas plants from more exposed sites can be rather tight.


The rather isolated position of this species is emphasized by its combination of flat leaves, 

stalked inflorescences and numerous seeds. In the molecular tree (Trift & al. 2004) it appears as 

sister group to Dionysia sect. Dionysia with moderate support, but except for the flat leaves there 

are no obvious morphological synapomorphies. 

VI. Dionysia sect. Dionysia 

= Dionysia subsect. Bryomorphae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 78. 

1961. – Type: D. bryoides Boiss. 

[– Dionysia sect. Epicamptophyllum Melchior in Mitt. Thüring. Bot. Vereins 50: 167. 1943], nom. 

inval. (Art. 22.2 ICBN, McNeill & al. 2006) as this name is based on the type of the generic name. 

[– Dionysia subsect. Caespitosae Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 77. 

1961], nom. inval. (Art. 22.2 ICBN, McNeill & al. 2006) as this taxon explicitly includes D. odora, 

the type of the generic name. 

31. Dionysia sawyeri (Watt) Wendelbo in Aarbok Univ. Mat.-Naturvitensk. Ser. 3: 64. 1961 ≡ 

Primula sawyeri Watt, Report Bot. Coll. SW Persia H. A. Sawyer: 94. 1891. – Holotype: W Iran, 

Kar Kanun, Kuhrang, Sawyer (not found by Wendelbo or me, probably lost). 

= Dionysia bachtiarica Bornm. & Alexeenko in Bull. Herb. Boissier, ser. 2, 4: 515, t. 2 fig. 3. 

1905 ≡ Primula bachtiarica Bornm. in Bull. Herb. Boissier, ser. 2, 4: 515. 1905. – Holotype: 

Iran, Kellar, Alexeenko 2722 (B, not seen, teste Wendelbo). 

The application of the epithet sawyeri is not absolutely certain, as the type is lost, but Dionysia 

bachtiarica from the same area, was considered synonymous by Wendelbo, based on the 

presence of stout marginal hairs on the leaves. I will accept this opinion. The type of D. bachtiarica 

is in B (Alexeenko 2722), but there are two more specimens in LE (Fig. 3) under different 

numbers (“ad rupes N jugi Kellar”, 8.9.1903 [cal. jul.], Alexeenko 821; “Vallis Lebze (?) in 

declivibus ad jugi Kellar ad rupes N”, 4.1902 [cal. jul.], Alexeenko 827). 

The colour of the corolla was given as yellowish with pale purple limb in the original description, 

which can be variously interpreted. The tube is often very pale in purple flowered species, 

but hardly yellowish. There are some purple flowered species with a yellow “eye”, and 

hybrids between yellow and purple species may show more yellow. Further, it is not uncommon 

for yellow corollas of Dionysia to turn brown, green or even greenish blue in herbarium specimens. 

The flower colour of this species is thus doubtful, although purple is perhaps the most 

probable. 

Dionysia sawyeri is only tentatively placed in the section Dionysia and was not included in 

the molecular study by Trift & al. (2004). It deviates in its glabrous corolla with deeply emarginate 

lobes and the high number of seeds per capsule. 24 ovules can be found in one ovary, compared 

to 3-5 in the other species. It differs also in the pronounced foliar heteromorphism. The 

early summer leaves (of “vegetative” shoots) are up to five times as long as subsequent (over-wintering) 

leaves, and much thinner and greener. The leaves are always completely glabrous 

abaxially, but have a varying amount of long hairs on the adaxial side (Fig. 3). 

The type of Dionysia sawyeri was collected by Sawyer in “Kar Kanun, Kuhrang”, whereas 

Alexeenko's collections are from “Kellar”. Kellar is, according to Wendelbo (1965), the same as 

Kuh-e Kukalar (= Kuh-e Kallar). Mozaffarian collected rich Dionysia material on Kuh-e Kallar S 

of Sibak, 4.7.1986, M 57420, 57399 (TARI). These plants, however, belong to D. lamingtonii. 

Zschummel looked for this species on the N side of Kuh-e Kallar in 2001, but found only D. 

caespitosa (Zschummel 01-12, UPS). Possibly, D. sawyeri does not grow in the same kind of habitats 

as most other species of Dionysia. 

32. Dionysia iranshahrii Wendelbo in Iran. J. Bot. 1: 72. 1976. – Holotype: W Iran, Bakhtiari, 

Kuh Pashmaku W Semirom, 6.1974, M. Iranshar (Ministry of Agriculture, Evin, Tehran; isotypes: 

GB, TARI). 

Like the previous species, this occupies a somewhat isolated position in the section, both in 

morphology and according to the molecular tree (Trift & al. 2004) where it is the sister species to


Fig. 3. Dionysia sawyeri – A: flowering branch with fresh axillary shoot; B: old shoot (overwintering); C: calyx; 

D: corolla; E: small leaf from above; F: marginal hairs of leaf. – From Alexeenko 821, LE. 

the rest of the section (Dionysia sawyeri not included). It has a glabrous corolla in common with 

D. sawyeri, but is otherwise not similar. 

In Grey-Wilson (1989) it is depicted with deeply divided calyx, but usually this is divided to 

only 1 /2 or at most to 2 /3. In the cliffs of the Kuh Pashmaku it is often found growing in places well 

protected from overhead water. It has proven rather difficult to keep in cultivation. 

33. Dionysia caespitosa (Duby) Boiss., Diagn. Pl. Orient. ser. 1, 7: 67. 1846 ≡ Gregoria caespitosa 

Duby in Candolle, Prodr. 8: 46. 1844 ≡ Macrosyphonia caespitosa (Duby) Duby in Mem. Soc. 

Phys. Genève 1844: 427. 1844 ≡ Primula macrosiphonia Kuntze, Rev. Gen. Pl. 2: 400. 1891. – 

Lectotype (designated by Grey-Wilson 1989): Iran, Elwend Kuh, near Esfahan, Aucher-Eloy 2609 

(K; isolectotypes: BM, P). 

= Dionysia peduncularis Bornm. in Bull. Herb. Boissier, ser. 2, 5: 261. 1905. – Holotype: W 

Iran, Kohrud, 4.1904, Strauss (B, not seen, teste Wendelbo). 

= Dionysia bolivari Pau in Trab. Mus. Ci. Nat., Ser. Bot. 14: 27. 1918 ≡ Dionysia caespitosa subsp. 

bolivarii (Pau) Grey-Wilson, Gen. Dionysia: 99. 1989. – Holotype: W Iran, Bazoft valley, Escalera 

1899 (MA, not seen, teste Wendelbo). 

According to Jamzad (1996) Dionysia bolivari is not distinct enough to be recognised as separate 

taxon. 

The corolla is glandular pubescent (illustrated with glabrous corolla in Grey-Wilson 1989) and 

sometimes has dense farina. Farina is rare in the section Dionysia, and is in D. caespitosa confined 

to the (upper part of the) exterior of the corolla and the interior of calyx. For the only other occasion 

of farina in this section, see discussion under D. diapensifolia.


34. Dionysia gaubae Bornm. in Feddes Repert. Spec. Nov. Regni Veg. 41: 179. 1937. – Holotype: 

Iran, Luristan, Khorramabad, Pole-Kalhor, 4.1936, Gauba (B). 

This rare species was recently refound (Lorestan: Peesh Kuh [32.58.26; 49.37.21], 2450 m. 

6.5 2002, T4Z 190, UPS). Its closest relative appears to be Dionysia odora, which differs in crispate-hairy 

leaves, more discrete leaf whorls and not strongly reflexed marcescent leaves. 

Dionysia gaubae var. megalantha Lidén, var. nov. 

Holotype: Iran, Lorestan: Peesh Kuh, T4Z 1028 (UPS) – Fig. 4. 

Caespites densiusculi efarinosi, caulis foliatus ramosus foliis marcescentibus valde revolutis 

obsitus. Folia viridia, obovata vel oblanceolata, hirsuta obsita, pilis longitudine varia glanduliferis 

praeter pilis longissimis eglandulosis. Flos plerumque solitarius, sessilis. Bracteae anguste 

oblanceolatae non vel parce dentatae, dense glanduloso-hirsutae. Calyx 8-10 mm longus, 

ad 4 /5 in lobis anguste oblongis fissus, extus et intus dense glanduloso-hirsutus. Corolla laete 

flava, 20-28 mm longa, saepe leviter curvata, extus dense glanduloso-hirsuta limbo 13-16 mm 

lato lobis ellipticis vel obovatis integris. In flore longistylo stylus ex corolla manifeste excertus. 

Ovarium 5-ovulatum. Capsula ignota. 

Etymology. – From Greek megalos = very big and anthos = flower. 

Fig. 4. A-B: Dionysia gaubae var. megalantha – flower (A); leaf, left adaxial view, right abaxial view (B); 

cutting from holotype cultivated in the Göteborg Botanic Garden. – C: D. odora – flowering plant; Prov. 

Kermanshah, Kuh-e-Parou, KMZ 9511, cultivated Göteborg Botanic Garden. – D: D. gaubae var. gaubae – 

flowering plant; T4Z 190, cultivated Göteborg Botanic Garden.


Suffruticose efarinose perennial forming rather lax cushions. Stems leafy and branched, densely 

glandular hairy when young, becoming pale reddish brown; new growth with more distant leaves 

at first, eventually very densely leafy towards the apex; dead (marcescent) leaves remaining on 

the stem for up to 4 years, usually rolled backwards, often as much as to form a little coil. Leaves 

green, 6-17 × 2.5-4 mm, gradually attenuate into a pale petiole again becoming slightly wider at 

the point of attachment, covered all over with glandular hairs, from very short or subsessile to at 

least 0.5 mm, and more sparsely with longer articulate hairs (up to 1.1 mm) that usually lack a 

gland; lamina obovate in outline, with 4-6 large blunt teeth; on each branch, the lower, first 

formed, sparsely set leaves are much longer than the ones in the apical overwintering flowering 

leaf rosette. Inflorescence sessile, 1(-2)-flowered. Bracts 1-2, 7-10 mm long, narrowly oblanceolate, 

entire or with a few teeth, densely glandular hairy on both sides. Calyx 8-10 mm long, 

cleft to 4 /5 into narrowly oblong lobes, glandular hairy on both sides. Corolla (long-styled) bright 

yellow (paler beneath), 20-28 mm long, often slightly curved, densely hairy with long and short 

glandular hairs; limb 13-16 mm broad, divided into elliptic to obovate slightly overlapping 

lobes. Style exserted 1-4 mm, eventually even up to 7 mm. Stamens situated very high up, 1 /4 to 

1 

/3 from the throat of the corolla. Capsule and seeds not known; number of ovules 5. 

This variety is similar to Dionysia gaubae sensu stricto in general habit and in the strongly 

back-rolled marcescent leaves, but differs in the much longer indumentum, the much larger calyx 

and the longer and comparatively narrower leaves. “Typical” D. gaubae has only glandular 

hairs, rarely longer than 0.2 mm. Our plant differs from D. odora in the much larger calyx, the 

much longer “vegetative” leaves that roll back when withering and in the large corolla with longer 

hairs on the tube. 

It grows in close proximity to the Dionysia gaubae population mentioned above (T4Z 190), and 

the specimen was selected for its large flowers. To emphasise its very deviating morphology I have 

chosen to give it a formal recognition but lack information on the variation pattern in the population. 

35. Dionysia odora Fenzl in Flora 26: 390. 1843 ≡ Primula odora (Fenzl) Kuntze, Revis. Gen. 

Pl. 2: 400. 1891. – Holotype: Iraq, Kurdistan, Mt Gara, Kotschy 386 (W; isotypes: B, BM, G, K). 

= Gregoria aucheri Duby in Candolle, Prodr. 8: 46. 1844 ≡ Dionysia aucheri (Duby) Boiss., Fl. 

Orient. 4: 19. 1879. – Holotype: Iran/Iraq, Nal Kuh, Aucher-Eloy 2832 (G; isotypes: BM, K). 

= Dionysia straussii Bornm. & Hausskn. in Bull. Herb. Boissier, ser 2, 3: 591, t. 6 fig. 1. 1903 ≡ 

Dionysia odora subsp. straussii (Bornm. & Hausskn.) Bornm. in Beih. Bot. Centralbl., Abt. 2, 

28: 462. 1911 ≡ Dionysia odora var. straussii (Bornm. & Hausskn.) Bornm. in Beih. Bot. Centralbl., 

Abt. 2, 33: 167. 1915. – Holotype: Iran, Kuh Gerru, near Burudjerd, 6.1902, T. Strauss 

(J(?); isotype: S). 

?= Dionysia odora var. integrifolia Bornm. in Beih. Bot. Centralbl., Abt. 2, 33: 167. 1915. – Holotype: 

not seen. 

= Dionysia sintenisii Bornm. in Bull. Herb. Boissier, ser. 2, 3: 592, t. 6 fig. 3. 1903. – Holotype: 

Turkey, Mardin, Bakahri, Sintenis 1282 (B; isotypes: G, K). 

36. Dionysia haussknechtii Bornm. & Strauss in Bull. Herb. Boissier, ser. 2, 4: 514, t. 2 fig. 2. 

1904 ≡ Primula haussknechtii (Bornm. & Strauss) Bornm. in Bull. Herb. Boissier, ser. 2, 4: 516. 

1904. – Holotype: Iran, Luristan, Shuturun Kuh, T. Strauss 1903 (B). 

37. Dionysia cristagalli Lidén, sp. nov. 

Holotype: Iran, Lorestan, 61 km from Aligodarz to Shoulabad, 2 400 m, lime cliffs, 29.6.1971. 

Runemark & Lazaro 26215 (TARI) – Fig. 5. 

Caespites densissimi grisei efarinosi, caulis numerosis foliis dense imbricatis. Folia anguste 

elliptica, subtus sparse et breviter glandulosa, supra et margine pilis glandulosis et eglandulosis 

obsita. Pili eglandulosi apicales basin confluentes cristam membranaceam albida formantes. 

Bracteae 1-2 linearo-oblongae foliis similes. Calyx usque ad basin in lobis anguste oblanceolatis 

fissus. Corolla flava, extus pubescens, lobis obcordatis. Ovarium 3-5-ovulatum.


Fig. 5. A-G: Dionysia cristagallii – bract (A); corolla (B); calyx (C); leaf, lower side (D); leaf, upper side (E); 

apex of leaf (F); flowering branch (G); from the type. – H: Dionysia haussknechtii – apex of leaf; from the type. 

Etymology. – From Latin crista =crestandgallus = cock. 

Dense cushions, rather similar to Dionysia haussknechtii, but more grey in appearance. Branches 

eventually becoming bare below, but densely clad with closely imbricate marcescent leaves for 

at least one cm, and more straight and “columnar” than those of haussknechtii; the difference between 

early and late leaves not pronounced, but the young overwintering shoots produced in autumn 

being quite discrete, ovate, with closely packed leaves. Leaves narrowly obovate-elliptic, 

3-4 × 1-1.5 mm, often slightly involute, at apex with a conspicuous crest of whitish basally confluent 

hairs; both sides of leaf beset with short-stalked capitate glands, longer hairs (both glandular 

and eglandular) confined to the distal part of the adaxial side (as is common in Dionysia). 

Flowers sessile. Bracts 1-2, like the leaves, but shorter and narrower, 2-2.5 × 0.5-0.7 mm. Calyx 

almost 3 mm long, divided to the base into narrowly oblanceolate lobes, similar to the bracts, and 

(like the leaves) equipped with an apical crest of basally confluent hairs. Corolla yellow, externally 

glandular-pubescent, (7-)9-12 mm; limb 5-6 mm broad, divided into obcordate emarginate 

lobes. Capsule narrower than in most other species of the genus, containing 3-5 seeds. 

The whitish membranous lacerate fringe that ornates the apices of leaves, bracts and calyx lobes, 

immediately set this species apart from all other Dionysia species. 

Additional records. – Lorestan, between Aligoudarz and Shoulabad, Ghadee Kuh (33.06.51; 

49.24.53), 2500 m, 5.5.2002, T4Z 175 (UPS); id., 7.5.2001, Zschummel 01-18 (UPS).


Fig. 6. Dionysia zetterlundii – A: bract; B: corolla; C: leaf, upper side; D: leaf, lower side; E: calyx; F: apex 

of leaf from above; G: flowering branch; H: hair details, left corolla, right leaf margin. – From plants cultivated 

in the Göteborg Botanic Garden from seeds of the type collection. 

38. Dionysia zetterlundii Lidén, sp. nov. 

Holotype: Bachtiari Charee pass to Bazoft valley (32.11.26; 50.11.58), 2700 m, 30.4.2002, T4Z 

125 (GB; isotypes: UPS, TARI) – Fig. 6. 

Caespites densissimi efarinosi, ramulis numerosis foliis dense imbricatis. Folia obovata obtusa 

glanduloso-hirsuta, superne cum pilis eglandulosis longis immixtis. Bracteae 1-2, foliis similes. 

Calyx ad 5 /6 in lobis oblongis fissus. Corolla flava, extus pubescens, lobi late obcordati. Ovarium 

4-5-ovulatum. 

Dense cushions, efarinose, with stems clad with marcescent leaves, eventually becoming bare in 

the older parts; most parts of the plant rather densely equipped with short to rather long glandular 

hairs. Leaves entire, obovate, obtuse to subobtuse, slightly involute, 3-4.5× 1.3-1.7 mm; in seedlings 

broad leaves with a couple of apical teeth may form, but soon succeeded by entire leaves; 

lamina densely set with glandular hairs of varying length, and on the adaxial side (especially in 

the distal part) with long eglandular straight hairs perpendicular to the leaf surface or (towards 

the upper margin) retrorse; rarely, long eglandular hairs occur also on the abaxial side of the 

leaves. Bracts 1-2, like the leaves, 3-3.5 × 1 mm. Calyx 3-3.5 mm, split to 5 /6 in oblong to oblong-elliptic 

obtuse lobes, glandular pubescent like the leaves. Corolla yellow, with 10-14 mm 

long tube, externally densely pubescent with short-stalked glands; limb 9-10 mm broad with 

broadly obcordate overlapping slightly emarginate lobes. Ovary with 4-5 ovules. 

Hardly to be confused with other species. Dionysia haussknechtii is superficially similar, but differs 

clearly in long marginal glandular hairs and absence of eglandular hairs, and in smaller calyx 

divided to the base. 

Growing on limestone, mostly on vertical and overhanging cliffs, in the Charee (Cheri) pass between 

the Kuhrang and Bazoft river valleys in West-Central Zagros. 

Eponymy. – Named after Henrik Zetterlund, a member of the T4Z team who found this species 

and brought it into cultivation. Henrik has – since the time of Per Wendelbo – maintained the


worlds largest living collection of Dionysia in the Göteborg Botanic Garden, a collection that has 

been and is of prime importance for research on taxonomy, phylogeny, anatomy and chemistry of 

the genus. 

Additional record. – Charee pass (32.09.44; 50.10.44), 3.5.2002, T4Z 140 (UPS). 

Note. – This species was provisionally called Dionysia heterotricha in cultivation in Gothenburg 

Botanic Garden, a name which – although never published - may have diffused into Dionysia circles. 

Due to the presence of the unrelated subsection Heterotrichae, this name would, however, 

cause confusion, and has therefore been abandoned. 

39. Dionysia tacamahaca Lidén, sp. nov. 

Holotype: Iran, Kermanshah, Bimar Mt., near Hukani Vill., South Kerend, 1500 m, Wendelbo & 

Assadi 16761 (TARI) – Fig. 7. 

Caespites densissimi, grisei, efarinosi, ramulis numerosis foliis marcescentibus tectis. Folia anguste 

obovata subacuta, dense glanduloso-hirsuta. Bracteae 2, lineares. Calyx ad 1 /2 usque ad 3 /5 

in lobis late ovato-lanceolatis subacutis fissus. Corolla flava, extus pubescens, lobis ovatis non 

emarginatis. Ovarium 3-5-ovulatum. 

Etymology. – Spanish tacamahaca (from Nahuatl: tecamaca) = balsam, with aromatic resin (cfr. 

Populus tecamaca = P. balsamifera). 

Dense aromatic greyish green cushions with stems in the upper part covered with marcescent 

leaves. No pronounced difference between early and late leaves. Leaves 3-4.5 × 1.3-1.6 mm, 

slightly bent outwards, narrowly obovate, subacute, densely covered on both surfaces with both 

short and rather long glandular hairs; the marginal hairs are longer in the lower half of the leaf; 

the longest hairs sometimes lack an apical gland. Flowers sessile. Bracts 1-2, linear, shorter than 

calyx. Calyx campanulate, 3.5-4.5 mm long, split about halfways into broadly ovate-lanceolate 

Fig. 7. Dionysia tacamahaca – A: corolla; B: calyx; C: bract; D: leaf from above; E: leaf in profile; F: hairs 

on leaf margin; G: flowering branch. – From the type.


acute lobes. Corolla yellow, externally glandular-pubescent; tube c. 10 mm long; limb 7-8 mm 

broad, lobes ovate-subacute. Ovary with 3 to 5 ovules. 

Dionysia tacamahaca cannot be easily confused with other species. It is slightly similar to D. 

zetterlundii, but differs in several characters, such as the rather homogenous indumentum, acute 

leaves, and the large calyx that is cleft only to half or at most up to two thirds. Tarakli & al. 2385 

differs from the type in larger calyx (4.5-5 mm) divided to two thirds into oblong lobes, but is 

similar in indumentum and leaf form (flowers are lacking). Possibly, the calyx lobes enlarge after 

flowering. 

Additional record.–Tarakli&al. 2835 (TARI). I have at present no locality information for this 

specimen. 

40. Dionysia lamingtonii Stapf in Bull. Misc. Inform. Kew 1913: 43. 1913. – Holotype: Iran, 

Bakhtiari, 7.1912, Lamington (K). 

This species has been collected several times in recent years by V. Mozaffarian (e.g., Kuh-e 

Kallar, Chebd, Chelgerd Kuh, Zardlich, all TARI). The small thickish blue-green eglandular 

leaves with distinct venation and long soft crispate hairs makes the species unmistakable. In earlier 

descriptions (Wendelbo 1961, Grey-Wilson 1989) the leaves are said to be glabrous above and 

hairy below. On the contrary, the long coarse hairs so typical of this species are more prominent on 

the adaxial side of the leaf. They may even be absent from the abaxial side, but never from the 

adaxial side. Pubescence varies a lot, even between successive leaves on the same stem. A segment 

with leaves glabrous on the outside can be followed by one with rather densely hairy leaves. 

41. Dionysia mozafarrianii Lidén in Iran. J. Bot. 8: 304. 2000. – Holotype: Chaharmahale Bakhtiari, 

Semirom, at the entrance into the city from the Shahreza road (3°26'N, 51°35'E), West-facing 

lime cliffs, 17.5.1998, Lidén, Mozafarrian, Popp & Seisums SLIZE 232 (TARI; isotype: 

UPS). 

42. Dionysia iranica Z. Jamzad in Iran. J. Bot. 7: 23. 1996. – Holotype: Bakhtiari, Lordegan, 

Monj, Badamestan Kuh, N of Bon-e-Gerd, 1500-2350 m, Mozafarrian 54700 (TARI). 

Recent material (T4Z 118, UPS) of this fine species, from more exposed sites at the type locality, 

deviates slightly from the type in that the plants are much more compact and the marcescent 

leaves not or only very slightly reflexed. 

43. Dionysia zagrica Grey-Wilson in Kew Bull. 29: 691. 1974. – Holotype: W Iran, Kuh-i-Sehquta, 

58 km N Pataweh (= Patureh), Hewer 2023 (K; isotypes: GB, W). 

This species has recently been recollected (Zhelgerd tunnel), probably close to the type locality. 

For a comparison with Dionysia khuzistanica, see that species. 

44. Dionysia khuzistanica Z. Jamzad in Iran. J. Bot. 7: 25. 1996. – Holotype: Khuzestan, Dehdez, 

Kuh-e-Sefid opposite Sar Sahra village and Bazoft valley, Mozafarrian 74001 (TARI). 

Very similar to Dionysia zagrica, and evidently closely related, but with considerably harder 

cushions and with shorter leaves. They both have rather stiff and thick flat leaves with the sclereid 

reinforcements clearly visible externally as more or less longitudinal striae, and yellow flowers 

with small narrowly elliptic usually entire lobes. The style in longistylous flowers is slightly 

exserted. The characters given by Jamzad (1996) to differentiate the two species do not hold (Table 

2.). She gave the corolla of D. khuzistanica as glabrous, in contrast to that of D. zagrica; actually 

both have hairy corollas, D. zagrica rather less so than D. khuzistanica. A similar situation 

obtains with the calyx. That of D. zagrica is if anything more deeply divided than that of D. 

khuzistanica, contrary to Jamzad. 

45. Dionysia termeana Wendelbo in Bot. Not. 123: 306, fig. 2g-i. 1970. – Holotype: Iran, Kuhi-Dena, 

Sisakht to Gadaneh-Bidjan, 6.1969, Termé 8131E (GB; isotype: TARI).


Table 2. Comparison between Dionysia zagrica and D. khuzistanica. 

D. zagrica D. khuzistanica 

Leaves obovate, subobtuse broadly ovate, acute, sometimes trifid 

Glandular hairs sessile very shortly stalked 

Calyx divided 4 /5-5 /6 divided 3 /4-4 /5 

Calyx lobes oblanceolate with narrow base narrowly triangular, broadest at the base 

Dionysia termeana is extremely variable in habit; plants growing in more shady situations are 

generally broad-leaved and lush, whereas plants from drier and more exposed stations are 

pulvinate with small leaves. It is thus understandable that it has been attributed to both “Bryomorphae” 

and “Caespitosae” by different authors. Experience from cultivation shows that this 

variation is to a large extent genetically based. It is erroneously illustrated by Grey-Wilson (1989) 

as having glabrous corolla. 

46. Dionysia michauxii (Duby) Boiss., Diagn. Pl. Orient., ser. 1, 7: 67. 1846 ≡ Gregoria michauxii 

Duby in Candolle, Prodr. 8: 46. 1844 ≡ Primula michauxii (Duby) Kuntze, Revis. Gen. 

Pl. 2: 400 1891. – Holotype: “Persia”, 1783-84, Michaux (G; isotypes: K, P). 

This species grows very abundantly on the mountain behind the university library in Shiraz, 

where it was collected by Wendelbo and Bokhari and later (1998) by the SLIZE expedition. As 

Michaux gave only “Persia” on his label, there is no reason to consider the other known locality 

“Bamu” to be the type locality, as stated by Grey-Wilson (1989: 124). The species has a peculiar 

dense pubescence of rather short retrorse acute eglandular hairs. It is not confined to vertical cliffs, 

but also grows on gentle slopes. 

47. Dionysia bryoides Boiss., Diagn. Pl. Orient., ser. 1, 7: 66. 1846 ≡ Dionysia kotschyi Bunge in 

Bull. Acad. Imp. Sci. Saint-Pétersbourg 16: 560. 1871, nom. illeg. ≡ Primula bryoides (Boiss.) 

Kuntze, Rev. Gen. Pl. 2: 400. 1891 [as bryodes] ≡ Primula kotschyi (Bunge) Kuntze, Rev. Gen. 

Pl. 2: 400. 1891, nom. illeg. – Holotype: Iran, Fars, Kuh Ayub, Kotschy G([?] ; isotypes: BM, K, 

UPS, W); probably same as Kotschy 406b: B, LE, P). 

One of the more widespread of the Zagros Dionysias, occurring as far north as the vicinity of 

Semirom (Gardane-e Shir, Nowroozi 2139, TARI). In general, the northern colonies seem to have 

slightly longer glandular hairs than those from Fars. 

48. Dionysia diapensiifolia Boiss., Diagn. Pl. Orient., ser. 1, 7: 65. 1846 ≡ Dionysia drabaefolia 

Bunge in Bull. Acad. Imp. Sci. Saint-Pétersbourg 16: 558. 1871, nom. illeg. ≡ Primula drabifolia 

(Bunge) Kuntze, Rev. Gen. Pl. 2: 400. 1891 ≡ Primula diapensiifolia (Boiss.) Kuntze, Rev. Gen. 

Pl. 2: 400. 1891, nom. illeg. – Holotype: Iran, Fars, near Persepolis, Kotschy 236 (G; isotypes: 

BG,BM,K,LE,PO,W). 

Dionysia drabaefolia was described from what appears to be an isotype of D. diapensiifolia 

in LE, distributed to P. 

This species may rarely have a peduncle up to 20 mm, and with 2 flowers. Some populations 

included here by Wendelbo, have entire leaves and corollas that may have farina on the apical outside, 

like is sometimes seen in Dionysia caespitosa. More research is needed. Possibly this represents 

an undescribed species. 

49. Dionysia sarvestanica Z. Jamzad & Grey-Wilson in Kew Bull. 44: 124. 1989. – Holotype: 

Iran, Fars, Post-i-Chenab SE Sarvestan, 6.1983, Mozaffarian 46754 (TARI; isotypes: K). 

A species not unlike Dionysia bryoides vegetatively, but the columnar stems are slightly 

broader, the glandular hairs on the leaves are slightly longer (i.e. still short, but never subsessile) 

and more dense, and the corolla is yellow with entire lobes and the style usually distinctly 

exserted in long-styled flowers. It is not impossible that some of the early records of D. bryoides 

from the mountains S of Shiraz could refer to this species, which grows on rather gently sloping 

rocks, as well as on more vertical sites.


a. subsp. sarvestanica 

b. Dionysia sarvestanica subsp. spathulata Lidén, subsp. nov. 

Holotype: Firozabad: Kuh-e- Sefida e-Meaymand, 24.3.1971, Dehgani 5466 (TARI). 

A subspecie sarvestanica foliis spatulatis valde latioribus facile distinguenda. 

Very dense cushions of closely packed cylindrical columns; leaves much broader and more 

rounded at apex than those of subsp. sarvestanica. 

Additional specimens examined. – Locus classicus: T4Z 1044 (UPS). 

Acknowledgement 

Several people have contributed to the new information presented in this article. First and foremost 

Valiollah Mozaffarian from the Institute of Forests and Rangelands, without whom much 

less material had been available for study. I also owe to his expertise and field knowledge all of 

the finds made during the SLIZE expedition 1989. Dieter Zschummel, Harry Jans, Michael Jaeger 

and Henrik Zetterlund have provided colour photos from wild stations, besides herbarium 

specimens, seeds, cuttings and observations on material in cultivation. Henrik Zetterlund and 

Gerben Tjerdsma keeps an outstanding collection of Dionysia in cultivation in Göteborg Botanic 

Garden, for which I am immensely grateful. The Tehran herbarium (TARI) generously put their 

very rich material at my disposal in 1998, and the staff at Komarov herbarium (LE) are heartily 

thanked for their hospitality. Finally, I wish to extend my sincere thanks to Hans Walter Lack 

and the editor, for their very accurate scrutiny of the first version of this paper, which has definitely 

increased its relevance. 

References 

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[CrossRef] 

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Biologia (Lahore) 34: 367-395. 

— & Wendelbo, P. 1976: Anatomy of Dionysia I. Foliar sclereids. – Notes Roy. Bot. Gard. Edinburgh 

35: 131-141. 

— & — 1985: Anatomy of Dionysia: II. Xeromorphic features. – Notes Roy. Bot. Gard. Edinburgh 

42: 327-345. 

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the Seventeenth International Botanical Congress Vienna, Austria, July 2005. – Regnum. 

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perspective. – Edinburgh J. Bot. 63: 21-48. [CrossRef] 

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by analysis of sequences from the chloroplast gene rbcL. –Syst.Bot.27: 396-407. 

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the subdivision of the genus. – Aarbok Univ. Bergen, Mat.-Naturvitensk. Ser. 1961(4). 

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Address of the author: 

Magnus Lidén, Uppsala University Botanic Garden, Villavägen 8, 75236 Uppsala, Sweden; 

e-mail: Magnus.Liden@botan.uu.se

The genus Dionysia (Primulaceae), a synopsis and five new species

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