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Physochlaina orientalis 04.jpg
Physochlaina orientalis
Scientific classification e
Kingdom: Plantae
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Solanales
Family: Solanaceae
Subfamily: Solanoideae
Tribe: Hyoscyameae
Genus: Physochlaina

6–10, see text

Physochlaina is a small genus of herbaceous perennial plants belonging to the nightshade family, Solanaceae,[1] found principally in the north-western provinces of China (and regions adjoining these in the Himalaya and Central Asia)[2][3] although one species occurs in Western Asia, while another is found as far east as those regions of Siberia abutting the eastern borders of Mongolia and also not only in Mongolia itself, but also the Chinese autonomous region of Inner Mongolia. The genus is a valuable one, since its species are not only of considerable medicinal value, being rich in tropane alkaloids, but also of ornamental value, three species having been grown for the purpose, although hitherto infrequently outside botanical gardens. Furthermore, the genus contains a species (P. physaloides – recorded in older literature under the synonyms Hyoscyamus physalodes, Hyoscyamus physaloides and Scopolia physaloides) formerly used as an entheogen in Siberia ( re. which see translation of Gmelin's account of such use below ).[4]

Derivation of genus name[edit]

The name Physochlaina is a compound of the Greek words φυσα ( phusa ), 'bladder' / 'bubble' / 'inflated thing' and χλαινα ( chlaina ), 'robe' / 'loose outer garment' / 'cloak' / 'wrapper' – giving the meaning 'clad loosely in a puffed-up bladder' – in reference to the calyces of the plants, which become enlarged and sometimes bladder-like in fruit – like those of the much better known Solanaceous genera Physalis, Withania and Nicandra, from which they differ in enclosing, not berries, but box-like pyxidial capsules, like those of Hyoscyamus ( see below ).[5] The variant spelling Physochlaena – as employed by Professor Eva Schönbeck-Temesy in her section on the Solanaceae for Flora Iranica – appears first on page 737 of Volume 22 of the German-language journal Linnaea for the year 1849.

Publication of genus name[edit]

The genus name Physochlaina was first published in 1838 by Scottish botanist George Don ( great-uncle of Monty Don ) on page 470 of volume IV of his four-volume work A General System of Gardening and Botany, often referred to as Gen. Hist. (an abbreviation of the alternative title A General History of the Dichlamydeous Plants) and written between 1832 and 1838. He included in his new genus the two species hitherto known as Hyoscyamus physaloides L. and Hyoscyamus orientalis M. Bieb. – the latter published by Baron Friedrich August Marschall von Bieberstein in his Flora taurico-caucasica of 1808.

Common names[edit]

Not being native to Western Europe, plants belonging to the genus Physochlaina have no common name of any antiquity in English, nor have they acquired a more recent common name among English-speaking gardeners, despite the passage of two centuries since their introduction to cultivation in the U.K.

Robert Sweet coined the English name Oriental Henbane for P. orientalis in his work The British Flower Garden in 1823, but this is simply a translation of the ( now obsolete ) name Hyoscyamus orientalis. He further coins the name Purple-flowered Henbane for the Siberian species P. physaloides, but this adds to the confusion, as, not only is the species in question no longer classified as a Henbane ( i.e. Hyoscyamus ) , but there are also a number of ( true ) Hyoscyamus spp. which bear purple flowers – e.g. Hyoscyamus muticus.

There is, however, a common name (age unknown) for Physochlaina in Russian, namely Пузырница (Puzeernitsa) – Bladder plant, qualified Пузырница Физалисовая (Puzeernitsa Phizalisovaya) – Physalis-like Bladder plant in the case of P. physaloides .[6] The Swedish common name for the genus – Vårbolmört – translates as 'Spring(-flowering) Henbane',[7] while the Finnish common name Kievarinyrtti means 'Inn Herb'[8] and the Estonian common name is Ida-vullrohu, meaning 'Eastern Henbane'.[9]

In the ancient, Iranian language Ossetian, spoken both to the North and the South of the Greater Caucasus range, plants of the genus Physochlaina have the common name Тыппыргæрдæг – approximate pronunciation Tippirgərdəg ( where schwa stands for the unique Ossetian vowel for which the special letter 'æ' had to be created in the Cyrillic alphabet ).[10][11] (See also page Physochlaina in Wikipedia, language: Ирон).

There are likewise several common names for the Himalayan Physochlaina praealta in the various languages of Nepal, and common names for the genus Physochlaina and the various Physochlaina species of Eastern Asiatic provenance in Standard Chinese (泡囊草属 pao nang cao shu), Tibetan (hun horse), Kazakh (үрмежеміс = (approximately) urmezhemis) Uyghur, Mongolian (garag chig tav) and certain Tungusic languages.[12]

Accepted species[edit]

The Plant List, a joint project of Royal Botanic Gardens, Kew and the Missouri Botanical Garden, accepts only six species of the genus:

The others being rejected mostly as synonyms.[16] Royal Botanic Gardens Kew Science Plants of the World online, however, accepts also:


Bisected fruiting calyx and separate operculum of Physochlaina physaloides
The yellowish-buff, pitted, reniform seeds of a Physochlaina species – probably P. physaloides, gathered in the Altai Mountains near the Mongolian city of Khovd in August 1989.

Perennial herbs, differing in their type of inflorescence – a terminal, cymose panicle or corymbose raceme – from the other five genera of subtribe Hyoscyaminae within tribe Hyoscyameae of the Solanaceae. Flowers pedunculate (not secund, sessile/subsessile as in Hyoscyamus). Calyx lobes subequal or unequal; corolla campanulate (bell-shaped) or infundibuliform (funnel-shaped), lobes subequal or sometimes unequal, imbricate in bud; stamens inserted at the middle of corolla tube; disk conspicuous; fruiting calyx lobes nonspinescent apically (i.e. lacking the spiny points characteristic of the calyces of the related genus Hyoscyamus – the Henbanes), fruiting calyx inflated, bladder-like or campanulate, loosely enclosing the capsular fruit. Fruit a pyxidium (i.e. dry capsule opening by a distinct operculum ( = lid ) – as in the other five genera of the Hyoscyaminae). Pollen grain polymorphic, usually subspheroidal, oval in polar view, circular-triangular in equatorial view.

Horticultural merit as ornamental[edit]

A gifted botanist blessed also with a gardener's eye for beauty, George Don is enthusiastic in his praise for the two plant species for which he created the new genus Physochlaina, noting in his ' A General History... ' of 1838 :

'The species of Physochlaina are extremely desirable plants; being early flowerers, and elegant when in blossom. They will grow in any soil, and are readily propagated by divisions of the root, or by seed. They are well adapted for decorating borders in early spring'.

In regard to the soil type favoured by wild populations, volume 22 of Linnaea (in surprisingly geological vein) provides the observation that Physochlaina orientalis is to be found growing on soils underlain by trachytes (volcanic rocks of a type notably rich in the chemical element potassium, a plant macronutrient essential for the production of flowers and fruit and, in a specifically Solanaceous context, the main ingredient of liquid feed for tomato plants).

Use in traditional Chinese medicine[edit]

Single flower of Physochlaina infundibularis Kuang – the 'hot ginseng' of Mount Hua, Shaanxi province.

At least three species of Physochlaina are currently used in traditional Chinese medicine : P. infundibularis, P. physaloides and P. praealta.

Physochlaina infundibularis[edit]

漏斗泡囊草 Lou-dou Pao-nang-ts'ao / lou dou pao nang cao (= 'Funnel-shaped Physochlaina '). The inhabitants of the neighbouring provinces of Shaanxi (rendered formerly 'Shensi') and Henan hold P. infundibularis in high esteem as a medicinal plant, regarding it as a kind of ginseng : most unusually for a toxic Solanaceous plant (totally unrelated botanically to the Araliaceous ginseng genus Panax) it is considered to be a 'general tonic' ( = adaptogen). The Chinese element 参 shen (= ginseng) forms a part of two of the common names for the plant, namely 华山参 Hua-shan-shen (= ginseng of Mount Hua) and Je-shen (= hot ginseng – from its hot, sweet, slightly bitter and astringent taste).

As with Panax, it is the fleshy root of Physochlaina infundibularis that forms the drug : the fresh, raw roots are first peeled and then boiled in a sugar solution containing small quantities of three other herbal drugs, before being dried, ready for storage and use. The three drugs added to the boiling solution are the root of Glycyrrhiza uralensis, the rhizome of Ophiopogon japonicus and the fruits of Gardenia jasminoides. This peeling, boiling and addition of 'cooling', 'yin' drugs is undertaken to mitigate the 'heat' / toxicity of the Physochlaina infundibularis roots.

In addition to its use as an adaptogen, P. infundibularis is used (in traditional Chinese medicine) in the treatment of asthma, chronic bronchitis, abdominal pain, palpitations and insomnia and as a sedative. The drug is also used to treat diarrhea of the kind considered in traditional Chinese medicine to be 'diarrhea due to deficiency of vital energy with symptoms of cold'.[20]

The nomenclatural association of P. infundibularis with Mount Hua – 'West Great Mountain' of the Five Great Mountains of China of Taoism – is an interesting one and merits further study : in common with other mountains regarded in China as numinous/Xian ling, Mount Hua (a precipitous assemblage of five (counted anciently only as three) peaks in the Qin range) is held to be a source of rare medicinal plants and life-prolonging elixirs. Furthermore, at the foot of the West Peak of Mount Hua (known as Lianhua Feng (蓮花峰) or Furong Feng (芙蓉峰), both meaning Lotus Flower Summit) stood, from as early as the second century BCE, a Taoist temple which was the site of shamanic practices undertaken by spirit mediums (see also Wu (shaman)) to contact an (unnamed) God of the Underworld and his minions, believed to dwell in the heart of the mountain.(See also Chinese folk religion).[21] Tropane-containing, Solanaceous plants (such as Datura and Hyoscyamus spp.) have a long history of use as entheogens in shamanic practices[22] – including Taoist practices[23]- and indeed Physochlaina physaloides is known definitely to have been used as an entheogen by certain Tungus tribes ( see section below ), so the possible use of its sister species P. infundibularis in Taoist, shamanic practices at Mount Hua might prove a topic worthy of consideration.

In addition to its being considered a kind of ginseng in its own right, the root of Physochlaina infundibularis ('Physochlainae Radix') is sometimes passed off in the ginseng trade as a substitute for the more costly roots of the true ginsengs Panax ginseng and Panax quinquefolius – a dangerous practice which could lead to the (potentially fatal), anticholinergic poisoning of unwitting users of these famous tonics, although the substitution tends to be a feature of local, Chinese (rather than international) trade.[24]

Physochlaina macrophylla[edit]

大叶泡囊草 Da-ye Pao-nang-t'sao / da ye pao nang cao. Like the root Physochlaina infundibularis, that of P. macrophylla has also (apparently) occasionally been passed off as that of Panax in the Chinese ginseng trade : '(The root of) Physochlaina macrophylla Bonati, a native of Honan, China, in appearance is very much like ginseng but slightly red; one should avoid using it as a substitute for ginseng as its alkaloid causes vomiting'.[25]

Physochlaina physaloides[edit]

泡囊草 Pao-nang-ts'ao / pao nang cao (= (common) Physochlaina) is the Standard Chinese name of the widespread species P. physaloides and the drug derived from it, which is used in the traditional medicine of Mongolia. In the traditional systems of medicine in China and Mongolia it is considered to have the effects of 'combatting weakness', 'warming up the stomach', 'soothing the mental condition' and relieving asthma. It is also used for treating 'diarrhea due to deficiency of vital energy with symptoms of cold' and 'cough or asthma caused by excessive phlegm or neurasthenia'. Note : the medical concept neurasthenia – now largely abandoned in Western medicine – is expressed in Chinese as shenjing shuairuo (simplified Chinese: 神经衰弱), a compound of shenjing 'nervous' and shuairuo 'weakness', and the Chinese condition so described is a culture-bound syndrome encompassing debility, emotional turmoil, excitement, tension-induced pain and sleep disturbances, caused by a depletion of qi ('vital energy') and impaired functioning of the wuzang (= 'five vital organs').[26][27]

Physochlaina praealta[edit]

西藏泡囊草 (H'si-Tsang Pao-nang-ts'ao / xi zang pao nang cao = 'Tibetan Physochlaina ' ) is the Standard Chinese name given both to Physochlaina praealta (Decne) Miers. and the drug prepared from its roots and aerial parts. This has been used in Tibet as a substitute for Tsang-ch'ieh (transliterated also as Zang Qie) – Anisodus tanguticus, more commonly known in China as shān làngdàng (= 山莨菪 = 'mountain henbane'). Unsurprisingly, for a tropane-containing plant, P. praealta has been recognised in India to have the belladonna-like property of causing mydriasis and is also used there as a topical medication in the treatment of boils.[28]

Use in traditional medicine of Tibet and Mongolia[edit]

Physochlaina species have a long history of use in the systems of traditional medicine of Tibet and Mongolia as drugs having powerful anti-inflammatory effects against skin diseases and sexually transmitted diseases, in addition to their beneficial effects – both soothing and energizing – upon nervous disorders. [29]

Hallucinogenic use of Physochlaina physaloides in Central Siberia[edit]

Johann Georg Gmelin, source of the first ( oft-quoted ) scientific account of the use of Physochlaina physaloides as an intoxicant in the Yenisei basin.
View of the Angara or Upper Tunguska River, a major tributary of the Yenisei – near the banks of which Gmelin encountered Physochlaina physaloides in the year 1738.
Pulmonaria officinalis : the herb for which a band of hungry Yeniseian Cossacks mistook Physochlaina physaloides – with dramatic results.
19th century depiction of hunting among the Evenks, one of a number of Tungusic peoples. Known in Gmelin's day simply as 'Tungus'. Makers and consumers of an hallucinogenic, Physochlaina-infused beer.
Ceremonial costume and drum of an Evenk shaman.Tropane-containing Solanaceae, such as Physochlaina spp., have frequently been employed in some forms of Shamanism to facilitate entry to an Altered state of consciousness.
Abbé Antoine François Prévost, author of the first 15 volumes of Histoire générale des Voyages, volume 18 of which made Gmelin's account of Physochlaina use better-known to subsequent scholars.
Professor Orazio Comes, author of the first work on recreational drugs to include Gmelin's account of Physochlaina use ( still under the incorrect designation 'Hyoscyamus' ).

Intrepid German naturalist, botanist and geographer Johann Georg Gmelin records in his Reise durch Sibirien of 1752 a remarkable account of the intoxicating properties of Physochlaina physaloides, which bears repetition in its entirety. On the 11th of August of the year 1738, Gmelin and his fellow explorer Stepan Krasheninnikov were negotiating the cataracts of the lower reaches of the Angara river – then known as the Upper Tunguska – in the Yenisei Basin, when they encountered a waterfall with a curious name :

...we came to Bessanova or Pyanovskaya D. which lies on the left bank of the river, and, two versts down, to another falls – Pyanoy Porog [ Russian : Пьаной Порог : 'The Drunken Rapids' ]...They were christened The Drunken Rapids by the first Yeniseian Cossacks to travel up from Yeniseisk on the stream and pass through them. They found in the vicinity of these rapids a herb, which they took, from the appearance of its leaves and flowers, to be Lungwort [ Russian : Медуница : Medunitsa ] and so used the leaves in the preparation of a vegetable soup and the roots to make a purée and, partaking of these dishes, grew so utterly intoxicated that they knew not what they were doing. When they had returned to their senses, they named these falls The Drunken Rapids and, because one suffers a headache after such a debauch, they named the falls that they encountered next Pokhmelnoy Porog [ Russian: Похмельной Порог : The Hungover Rapids ].

His curiosity aroused, Gmelin investigated, and discovered an attractive new species :

This account has given me the opportunity to reveal the identity of the beautiful plant involved, which was unknown to any botanist before me : Hyoscyamus foliis integerrimis calicibus inflatis subglobosis [ Botanical Latin : 'The Henbane having simple, untoothed leaves and ( fruiting ) calyces that are more or less round and inflated' [ i.e. like those of a Physalis ] Linn. h. Ups. 44. 2.

Having identified the ( Linnaean ) genus Hyoscyamus to which the intoxicating plant of The Drunken Rapids ( since moved by Don to the genus Physochlaina ) belonged, Gmelin went on to quiz his local guides and learned the following concerning its intentional consumption :

If one steeps the leaves or even the finely-chopped roots of this plant in brewed beer – or, better yet, in beer that is still undergoing fermentation – then it takes but a single glass of such beer to make a man exceedingly foolish : it is surely a strange draught that he quaffs, for he is robbed of all his senses, or at least finds his senses grossly disordered, mistaking tiny things for huge ones : a straw for the thickest of beams, a drop of water for a mighty ocean and a mouse for an elephant. Wherever he goes he encounters [ what he imagines to be ] insurmountable obstacles. He pictures continually to himself the cruellest and most dreadful imaginings of an inevitable death awaiting him, and, as it seems, all this fills him with despair, because his senses are withering away; thus, should one such drunkard go to step over a beam, he will take a great stride out of all proportion to the actual size of it, while another will see deep water in front of him [ where there is only shallow ] such that he dare not venture into it.

In conclusion, Gmelin then adds, concerning the plant itself :

The local inhabitants often use these roots when they want to play a prank upon each other. The Russian merchants often bring these roots back with them when they return to Russia, because they maintain them to be a sovereign remedy for bleeding haemorrhoids and also against the haematuria – a claim which I have been unable to verify.

[30] Gmelin's Reise durch Sibirien – with its evocative account of his findings concerning the plant now known to science as Physochlaina physaloides – received a translation into French which was published as part of Volume 18 of Abbé Prévost's monumental Histoire générale des voyages – a compendium of eighteenth century exploration by land and sea, continued beyond the original fifteen volumes, by other authors following the death of Prévost in 1763. The Histoire translation is by no means always a word-for-word rendering of Gmelin's original text, and, in the passage concerning Physochlaina, a sentence entirely absent from the Gmelin account has been added, which nonetheless has been retained in subsequent retellings of the passage in question :

Il parle continuellement sans savoir ce qu'il dit. [Translation : 'He speaks continually, without knowing what he is saying' – said of the man intoxicated by a single glass of potent Physochlaina beer].


The first work devoted exclusively to recreational drugs to draw on Prévost's translation of Gmelin's account of Evenki Physochlaina use was A History of Tobacco with notes on the use of all Excitants currently known by Italian botanist Professor Orazio Comes, written in French and published in Naples in 1900.[32]

Comes's summary of the Prévost translation was included by German Botanist Carl Hartwich in his classic and influential work of 1911 Die Menschlichen Genussmittel ( = 'The Pleasure-drugs of Mankind' )[33], which, in turn, was quoted by 21st century expert on hallucinogens Dr. Christian Rätsch in his Encyclopedia of Psychoactive Plants of 2005. Hartwich speaks only of 'Hyoscyamus' with no indication of the species involved and, while Rätsch uses the correct species name physaloides he still includes the plant in his discussion of the various Hyoscyamus species – seemingly unaware that the plant was actually made the type species of the new genus Physochlaina by George Don as far back as the year 1838.


Physochlaina species have yielded a variety of tropane alkaloids, including not only the hyoscyamine and scopolamine present also in better-known Solanaceous genera such as Atropa, Hyoscyamus and Scopolia, but also the new (eponymous) base physochlaine, first isolated from the aerial parts of the Central Asian species Physochlaina alaica Korotkova ex Kovalevsk.[34] Other tropanes present include apoatropine, aposcopolamine and 6-hydroxyatropine.[35]

Westernmost species : P. orientalis[edit]

Flowers of Physochlaina orientalis in profile showing pubescent calyces.
Capitate inflorescence of Physochlaina orientalis in profile.
Physochlaina orientalis flowers giving way to immature fruits.
The ripe, fruiting calyces of Physochlaina orientalis (M.Bieb.)G.Don, (length circa 18mm) showing nodding growth habit and indumentum of sticky trichomes with trapped insect remains and thistle pappus.
View of interior of ripe, fruiting calyx of Physochlaina orientalis (M.Bieb.) G.Don showing pyxidial capsule with detached operculum revealing yellowish-buff, pitted, reniform seeds.

Habitat in cryptis circa acidulam Narzana et in Iberia. Floret primo vere. – Marschall von Bieberstein. Flora Taurico-caucasica 1808

Confusingly, the species of Physochlaina most commonly encountered in cultivation not only bears what appears to be a counter-intuitive specific name, but is also no longer an accepted species : the plant grown as an ornamental under the name Physochlaina orientalis (M.Bieb.) G.Don, far from being ( as its specific name appears to imply ) the Physochlaina species with the easternmost distribution is, in fact, that with the westernmost, as it is native to eastern Turkey, southern Russia the Caucasus and north-western Iran.[36][37]

This apparent misnomer is an artifact of the plant's having initially been placed in the henbane genus Hyoscyamus as H. orientalis before the creation of the genus Physochlaina and the discovery and naming of its (Physochlaina 's) species of predominantly Chinese provenance.

The plant cultivated under the name Physochlaina orientalis (referable possibly to P. physaloides – see below) is a rhizomatous, clump-forming, perennial, up to 45 cm in height, bearing attractive, funnel-shaped flowers of a pale purplish-blue, followed, in fruit, by pubescent calyces much longer than the capsules enclosed.

In cultivation in the United Kingdom it can flower between March and May, flowering usually in the month of April, when it can make a fitting companion for Spring-flowering bulbs, particularly those sharing its preference for well-drained soil – indeed its Summer dormancy (an adaptation to drought, characteristic of Mediterranean vegetation (- see also Dry season)) resembles that of many genera of bulbous plants e.g. Tulipa.[38]

Despite its merits as a garden flower, P. orientalis is still seldom to be seen in British gardens, although it has been grown in Britain since at least 1818 – as noted by Robert Sweet :

This pretty Spring-flowering plant was raised from seed, received from Moscow, by Messrs. Whitley, Brames and Milne, at Fulham in the year 1818.


[ Note: the Fulham nursery of the above-mentioned Whitley, Brames and Milne was founded originally by Matthew Burchell ( c. 1752–1828 ),[40] father of the celebrated naturalist William Burchell. It was owned subsequently – in various partnerships – by nurseryman Reginald Whitley ( c.1754–1835 ).[41]]

In the wild, near the historic, Turkish, silver-mining town of Gümüşhane (on the westernmost edge of its range) P. orientalis is frequently to be found growing near cave mouths and in rock crevices[42]- exactly the type of microclimate referenced by Marschall von Bieberstein in his original description of 1808, where he speaks of ' grottos near the acidic mineral springs of Narzana (= Narzan Baths, Kislovodsk, North Caucasus) '. ( Compare also a similar penchant for growing in rock crevices on the part of the Xinjiang species Physochlaina capitata – see above ).

The plant's country of origin is given in von Bieberstein's original description of ' Hyoscyamus orientalis ' (now Physochlaina orientalis) as Caucasian Iberia – a former kingdom, the heartland of which is the modern Georgian province of Kartli. The Caucasian Kingdom of Iberia also encompassed parts of Armenia, Azerbaijan, southern Russia and eastern Turkey.

Flora Iranica is in agreement on this range of occurrence for P. orientalis, adding also to the list of territories not only north-western Iran but also 'Syr Darja' – the latter being referable to lands traversed by the river Syr Darya and, more especially the historic Syr-Darya Oblast and hence modern Uzbekistan. Uzbekistan lies outside the area encompassed by Flora Iranica, but parts of neighbouring Turkmenistan do not. Either way, Flora Iranica is unequivocal in describing the range of Physochlaina orientalis as extending eastward into Central Asia. In this context, it may be noted that Phillips and Rix include in their work on garden perennials a photograph of a second, unaccepted Physochlaina species of unequivocally Central Asian provenance, namely P. alaica Korotk. ex Kovalevsk, recorded as growing in the Pamir-Alay, a Central Asian mountain range taking in parts not only of Uzbekistan, but also of Tajikistan and Kyrgyzstan. Physochlaina alaica differs from P. orientalis in having flowers of a beige or yellowish-buff colour and corolla 'throats' veined within in a contrasting dark brown. As is to be expected of a plant native to the Pamirs, this species is extremely hardy, if grown in dry (i.e. well-drained) conditions.

Physochlaina physaloides and P. orientalis[edit]

Physochlaina physaloides in Curtis's botanical magazine (No. 852) (8469918743)
Physochlaina sp. (exserted pistil and stamens suggest P. orientalis, rather than P. physaloides). Afbeeldingen van zeldzaame gewassen (Tab. V) BHL285489

George Don notes of Physochlaina orientalis in his ' A General History... ' entry on his new genus :

This is very like P. physaloides; but differs in the higher stature, and more robust habit; in the herb being pale green, and more downy; the calyx being longer; and in the tube of the corolla widening gradually to the top; in the genitals being usually exserted; and in the calyx being less inflated, and hardly twice as long as the capsule.

Height, robustness and also, to an extent, stem and foliage colour being omitted from the discussion as functions of genetic strain, habitat and nutrition, one is left with relative pubescence, flower shape, exsertion of style and stamens and length and degree of inflation of the fruiting calyx as means of differentiating Don's original two species.

If Physochlaina orientalis were to be demoted to a subspecies of P. physaloides, one would be left with a single, rather variable species, found over an immense range stretching thousands of kilometers from Eastern Turkey through Iran, Central Asia, China and Mongolia all the way to southeastern Siberia.

Given the Central Asiatic provenance of the not-universally-accepted species Physochlaina alaica and P. semenowii and the assertion in Flora Iranica that P. orientalis may be found in Central Asia, it may be that more than one Physochlaina species will be subsumed in the concept of a variable and very wide-ranging P. physaloides.

Such variability and wide distribution bear comparison with those of a much better-known Solanaceous plant : Atropa belladonna, which a consultation of the literature will reveal to have acquired a relatively large number of specific and subspecific names now largely reduced to synonymy with A. belladonna as local varieties of a single very variable species found from the U.K. in the West to northern Iran in the East.[43]


Shoot development and anthesis in Physochlaina orientalis


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  8. ^ Google translate: Finnish to English.
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