Sunday, 20 December 2020

Shindal Makudi - Part Two

Continued from Part One 

Cultivation in India

One of my two plants, again. Growing in Manchester.

We don't know whether Shindal Makudi was cultivated deliberately in the region where it now occurs. It is possible that the plant was deliberately introduced to some of the hill forts. It is odd that more than half of the locations where it grows have been associated with old military forts or ancient Buddhist monasteries. A plant that provided both medicine for wounds and an edible vegetable would seem to be useful to have around in a military fort.

Though it is true that many of the high hills in Maharashtra have old hill forts on them. 

I have given details among the reports of collections from the wild in the first part of this article for each cultivation of which I have found reports. We know that the plant has been cultivated in Maharashtra state from collections by botanists from 1897, 1938, 1950 and one before 1971. The plant was also recorded as cultivated in northern central India, at Jaipur in Rajasthan in 1972 and Delhi in 1982. The origin of those plants was not recorded.¹⁵ ⁶⁴

The succulent enthusiast Jagdash Singh Sarkaria saw Frerea still being grown in December 1975 at the Western Circle headquarters of the Botanical Survey of India at Pune/Poona. Sarkaria was there consulting⁹³ the expertise of a Dr K Singh and the author of a short note on cultivated Frerea written in September 1975, RS Raghavan.⁷⁸

With several collections by Sarkaria⁹⁴ in the mid-1970s and so many others since, we can assume the plant has been in continuous cultivation in India since then, in botanical gardens⁵⁹ and private collections. 

I have, unfortunately, not found anywhere near as much detail on Indian cultivation as I have on European succulent growers.

Cultivation in Europe

Gordon Rowley thought that the plant was first seen in a living collection in Europe in 1958.⁸⁴ He encountered five rooted cuttings⁸⁵ in the greenhouses of L. Maurice Mason of Talbot Manor in Fincham in Norfolk in the east of England on the 24th of May 1958. Mason was a rich farmer who travelled the world to add to his vast collection of tropical and subtropical plants. Mason was famously generous and shared the cuttings with his fellow enthusiast. 

Obviously, decentralised cultivation of rarities increases the likelihood that the plants will survive under the care of somebody and then the experience and cuttings might be returned to the original donor if they had lost the plant. Or, as Maurice Morris said⁸⁴ "If you want to keep a plant, give it away!" 

The illustration of the plant was a photo of a pair of small cuttings with leaves but no flowers. Rowley did not see flowers until later in the year.⁸⁵ Rowley described Frerea, unfortunately ambiguously, as "introduced from India by Mr. Mason for the first time". So we do not know from this statement if Mason collected it himself or was sent the plants from connections made in India.

Four other recipients of Mason's generosity were Ron Ginns, Rowland (C.R.) Tyrrell, John Measures and Reg Collis, who were all also members of the National Cactus and Succulent Society.²⁸ ¹⁰²

In his 1974 recollections in a short piece in Asclepiadaceae, Ginns wrote²⁸ of visiting Mason, with Measures and Rowley: 
Mr. Mason is one of the greatest plantsmen of the present day and his extensive grounds and numerous greenhouses form a veritable botanic garden. We were privileged to be taken into his "holy of holies", a relatively small house crammed with rarities received from collectors all over the world.We were shown with pride a large pan of FREREA in flower.
This must have been later in 1958. Frerea will grow and flower rapidly from cuttings in the right conditions.

On his first visit to Mason, Tyrrell was embarrassed to be given over 300 assorted plants and cuttings. Tyrrell visited Mason again 18 months after first seeing a "pan of Frerea" at Talbot Manor. Mason remembered promising a Frerea and mentioned it as soon as Tyrrell arrived.¹⁰² If Tyrrell's "some 18 years ago" was correct, that first visit would have been in 1956. I assume this was fuzzy recall when the article was written and then published in 1974. I think Tyrrell must have received his plants in 1960 at the earliest.

Within the year Tyrrell persuaded his new plants to produce seeds and he raised eight seedlings the next year.¹⁰² Tyrrell was well-known (and often teased, especially by Ron Ginns) for watering and feeding his cacti and succulents far more heavily than other members of the NCSS.⁴⁸ He even signed himself C.R. (Lush) Tyrrell in a letter published¹⁰¹ in the The National Cactus and Succulent Journal defending the growing of "a green jungle of lush stems". Perhaps the cow manure he used also attracted a few of the right sort of flies for pollination of stapeliads.

Ginns wrote²⁸ (in 1974) of their visit to Mason: 
It is from these that most of the plants of FREREA in cultivation in this country are descended.
I think EW Putnam, who wrote the obituary for Ron Ginns in The National Cactus and Succulent Journal, may have misconstrued this sentence. I would assume that Ginns was referring to the large pan of Frerea in Mason's greenhouse as the origin of most British-grown Frerea. Putnam wrote⁷⁷ that 
He was probably correct in believing that all the plants of Frerea indica in this country derived from a plant of his.
Ginns was renowned for his generosity and skill at propagation of cacti and succulents. However, despite teasing Tyrrell about pampering his cacti, Ginns gave credit²⁸ to Tyrrell for his success with sub-tropical species like epiphytic cacti with his heated greenhouse in the tiny village of Welford on the border of Northamptonshire and Leicestershire in the English Midlands. Ginns wrote:
This treatment suited FREREA which was extensively propagated are [sic] distributed amongst the many visitors to Welford.
In 1982, Brian Fearn visited Maharashtra to collect succulents. He was particularly interested in finding Frerea indica. Fearn wrote about the information he was told by Professor PV Bole of St Xavier's College, Bombay:
Professor Bole has had a long interest in Frerea and reintroduced it to Western cultivation in 1960, sending plants to Dr. Hardy in S. Africa and to Maurice Mason in England. Maurice Mason later told  Professor Bole when he subsequently visited England that he had lost the plants. This clone was collected from Purandhar. Mr Gogari of Ahmadabad [sic] also collected plants from a site near Pune, which were sent to Edgar Lamb, who also sent some to Maurice Mason. This clone has subsequently been widely distributed. It is now believed to be extinct in the area around Pune itself, from where it was originally collected by Mr. Gogari, a chemical factory having been built on the site.
This passage is helpful and frustrating at the same time. Perhaps, after over 20 years Professor Bole was vague about the exact date that he had sent the plants to David Spencer Hardy (see below) and Maurice Mason. 

Clearly they are referring to Parvati Hill in Pune, though there are no chemical factories built on the hill. The only buildings appear to be temples. That had perhaps also altered in their memory from the phrase used by Sarkaria: "several years ago, when Poona had not become the big industrial city that it is today."

In a brief comment⁸⁶ under Fearn's article, Gordon Rowley wrote:
My slides (Figures 2A & 2B, page 25) are of the first (of 3?) Caralluma frerei plants at Talbot Manor. As you see, they pre-date the 1960 and later accessions. This is the original dark-flowered clone, the same that was illustrated in black and white in Nat. Cact.Succ.J. 13 (3): 47, 1958.
It looks as though Maurice's guests were rather too free to accept his advive [sic] "Help yourself!" - he had to replace his stock of the species three times.
From this, it seems Rowley took the 1960 date given by Bole as exact. The slides that Rowley mentioned show a flowering plant and a close-up of a flower. The pattern on those flowers seems to be of the same type as my plants and those of Edgar Lamb from 1961,⁴⁴ David Spencer Hardy³⁰ from 1963 and those photographed by Keith Grantham¹⁵ at Purandhar Hill in 1992.

I have not found any published comments from Edgar Lamb nor David Spencer Hardy on the origin of their plants.

Rowley also referred to the introduction of the plant to Norfolk very briefly in his A History of Succulents, published in 1997. In a summary⁸⁷ of Mason's life he wrote:
Larger than life champion of tropical plants at Talbot Manor in temperate Norfolk, whose glasshouses and garden stocked with exotics from his travels drew gardeners, botanists and specialists from far and wide, few of whom ever left empty-handed. He introduced Frerea indica (Caralluma frerei), some Peperomia species and other succulents to cultivation, and was President of the National Cactus & Succulent Society 1962-1969.

There is a photo of a flowering Frerea indica credited to the "Division of Botany, Pretoria" published³⁰ by the South African plant collector David Spencer Hardy³⁸ in November 1963. The name Division of Botany suggests that the photo was from between 1958 when Hardy started there to 1961 when it changed its name to the Botanical Research Institute. So, that does not help pin down the exact date that the plants were sent from St Xavier's College.

I did email both the Herbaria of the Pretoria National Botanical Garden and St Xavier's College, Mumbai to ask about this. This is not a good year to catch people in the office, I am afraid.

According to David J Goyder, a specialist in the Apocynaceae at Kew Gardens, in response to my question: 
the earliest cultivated material at Kew dates from 1959, when a plant was donated by L.M. Mason. The Kew accession number was 1959-65112, and a portion of the plant was preserved at the same time in Kew’s spirit collection⁵⁴ under the collection number Mason 651...

Les Cèdres is a huge private botanic garden in Saint-Jean-Cap-Ferrat in the Alpes Maritimes department of France on the Mediterranean coast, close to Monaco and the Italian border.

Les Cèdres was run from 1924 to 2016 by the family that made Grand Marnier liqueur. Julien Marnier-Lapostolle started developing the exotic plant cultivation there in 1928. In the issue of Cactus for October/December 1960, Marnier-Lapostolle commented that they had just received a Frerea indica.⁴⁹ There were no other details except that the habitat was near Pune. There is a photo of the flower, simply credited "Photo Rauh."

The comment "near Pune" might seem to favour an origin from Mr Ghoghari. However, Purandhar Hill Fort is only about 27km (17 miles) from Parvati Hill in the middle of the city of Pune.

Werner Rauh of the University of Heidelberg remarked⁸¹ in 1961 that it was strange that the plant was found in few European collections despite being easy to grow. Rauh's article is illustrated with two photos of the plant. One is in leaf but not flower, the credit is "Phot. W. Rauh". We might assume this is Rauh's own plant. However, the other photo is a close-up of a flower labelled "Frerea indica, Einzelblüte [single flower]. Jardin Botanique „Les Cèdres" Cap Ferrat Phot. W. Rauh". 

Although I have found no record of the travels of this particular plant, it is clear that all the European growers who had it in their collections knew each other. In 1966, Werner Rauh, Julien Marnier-Lapostolle and Maurice Mason were all founding members of the African Succulent Plant Society, of which Gordon Rowley was President.³ David Spencer Hardy and Werner Rauh had collected plants together. I can find no reference to either of them visiting India. 

The University of Heidelberg website does not list Frerea among its living collections. Their only online herbarium specimen is spirit-pickled and has the collector and donor as unknown with the date as 27/09/1972.  They have two specimens of the hybrid Frerea indica x Caralluma europaea without any information at all.

On the 11th of September 1962, Mrs M Stillwell gave a lecture⁹⁷ to the Cactus and Succulent Society of Great Britain at the RHS Lindley Hall Restaurant in Westminster in Central London. The talk was about observations and cultivation tips for succulents. Among her favourite species of stapeliads she mentioned:
Frerea indica has whitish stems with thick leathery leaves and flowers like a Caralluma.

Frerea indica was first flowered in the greenhouses of the Muséum National de Histoire Naturelle in Paris sometime in either 1964 or 1965. They gave no further details.⁸³ 

In 1966, in an article in the Cactus and Succulent Journal of Great Britain, Miss Margaret J Martin mentioned Frerea, illustrated with her photo of a leafy plant without flowers. The only photo in the article not credited to her was one of Stapelia flavirostris, credited to E. Lamb - presumably Edgar Lamb.⁵²

In 1967, Jürgen Reiss of Bad Kreuznach in West Germany wrote⁸² that he had received a specimen "some time ago" (Vor einiger Zeit...) which he successfully cultivated and brought to flower. There are two photographs to illustrate the article. However, the second photo appears to be the first one cropped to the flower and rotated anti-clockwise by 90º.

L. Maurice Mason

I feel that at this point I should get a little distracted and describe the enthusiastic plant hunter and grower, L. Maurice Mason.

Mason often exhibited his plants and won many awards at flower shows for begonias, bromeliads, orchids and more. Mason achieved at least 190 awards from exhibiting at the Royal Horticultural Society shows.  He was given the Royal Horticultural Society Victoria Medal of Honour, which could only be held by 63 people at any one time. Mason was a member of various committees of the Royal Horticultural Society from the 1950s to the 1980s. In 1957, the Agricultural Research Service of the United States Department of Agriculture credited Mason with 225 ornamental introductions to the USA.⁵⁷ 

Mason's friend Christopher Lloyd wrote⁴⁷ in The Guardian Weekend newspaper supplement in 2003:
Maurice was also a great character and bon viveur. His hospitality was prodigious and he was immensely generous. If you expressed pleasure in any plant seen as he took you round his garden, he would say, "You like?" and note it down, and presently you would receive a large parcel of all these goodies.
Mason was not appreciated by everyone. Especially later in his life when he was on orchid committees. Many of the the younger orchid growers found him to be typical of the "old fraud" orchid judges. Having encountered Mason in 1981, Andy Easton wrote:²²
The chairman of the committee was a gin-ridden old bombast by the name of Maurice Mason VMH. Pompous and condescending to all and sundry and typical of that British class who inherit and do little to make the world a better place.
One opinion²³ of him, from a secretary of the RHS (when asked about horticulturally-inclined dukes) was that Mason "acts like a Duke, though he ain't."

The most famous of Mason's many introductions to Britain was probably the Iron Cross Begonia that is still grown as a houseplant. This strikingly unusual plant is loved by some but hated by others. Mason was given the living plant when visiting Singapore in 1952. Mason arrived in Singapore on the 18th of March 1952 and stayed for three weeks. On the 28th of March 1952 he and his wife Margaret were judges at the Singapore Flower Show.⁴ 

The Singapore Free Press reported:⁵
Some of the Rex Begonias, Singaporeans will see for the first time at today's Flower show are a gift from Mr. Mason to some amateur growers in Singapore.
Having satisfied himself that his last gift of plants are thriving, he has promised to send more.
Much to the delight of growers, he has found a number of interesting plants here which he will take away with him.
In 1959, the Iron Cross Begonia was determined to be a species rather than a hybrid and named Begonia masoniana by Edgar Irmscher. Rudolf Christian Ziesenhenne considered the name to be improperly published because a type specimen was not designated and re-published it in 1971 in the same journal, The Begonian. Both used cultivated material to describe the species. 

Begonia masoniana was finally found in the wild on the border of Guangxi (in southeast China) and Vietnam in 2003 and 2004. The plants were found to be frequent on limestone rocks on the very edge of the city of Pingxiang. In 2019, more plants were found at two more locations, 35km (22 miles) and 45km (28 miles) to the north of Pinxiang.⁷

Mason did not just rely on his many connections with other gardeners. During their many travels during the winter months when the farm was less demanding, Mason and his wife Margaret would often venture into the wilds to find new species. They collected from Borneo, British Guiana, Costa Rica, Madagascar (three times), Malaya, New Guinea, South Africa and Tanganyika in just a few years in the 1950s.⁵⁷ They also collected from Australia, New Zealand, the Congo, Ghana, Brazil, Colombia and the Andes.²³

Brent Elliot wrote²³ of Mason in The Plantsman:
If he maintained detailed records about his finds, he did not publish them, so it has not usually been possible for subsequent botanists to determine whether he collected a particular plant in the wild, found it in cultivation, or received it as a gift.
An example of this problem was Sansevieria masoniana, a relative of the common houseplant, Mother-in-Law's Tongue. The botanist who named it after Mason was unable to determine whether Mason had collected it himself.

L Maurice Mason certainly left British horticulture quite changed, from his many exotic plant introductions, spectacular exhibitions and tireless promotion of the art of gardening.

Speculation on the origin of the European plants

Most of the pictures of flowers of Frerea from 1958 to 1967 in Europe have the distinctive appearance of the flowers of my plants, with their five letter-like glyphs. This clone is still commonly available in Europe. So, from where did Mason obtain his cuttings?

Though there is no precise evidence, there are some circumstantial indications that we might use to determine the origin of the European cultivar of Frerea indica.
  • The cuttings Mason received by May 28th were in full leaf and growth.⁸⁴
  • The mature sections of those cuttings do not seem to have the rugged silveriness of a plant that has endured a dry season.
  • The monsoon season in Junnar is from June to October, though there is some rain in May.
  • Mason preferred plant-hunting in winter, when there was less to do on the farm.²³
  • Hermenegild Santapau published accounts of his cultivation of Frerea at St Xavier's College in Mumbai in 1950⁹⁰ and 1957.⁹¹
  • The drawing of the plant collected at Vajragad by Santapau and Irani shows those five distinctive glyphs in the middle of the petals.⁹²
As the cuttings arrived during the dry season in Maharashtra, clearly the cuttings were from cultivated plants that were artificially watered. 

Both the 1950 and 1957 publications by Santapau can be found in the Library at the Royal Botanic Gardens at Kew. Mason also had an extensive botanical library of his own. Santapau and his student Irani made repeated collections of the plant between 1950 and 1960.

Though there were two reports of cultivation of Frerea indica in India before this, I think it unlikely that Mason would have tried to obtain those. The first was at least 55 years before and the second 20 years before Mason received his cuttings.

Was PV Bole out by two years when he told Brian Fearn in 1982 that they had sent plants to Mason in 1960? Just as Rowland Tyrrell was out by two years when he recalled visiting Mason?

PV Bole was born and brought up in Saurashtra in Gujarat State. He became Head of the Botany Department of St Xavier's College and Director of the Blatter Herbarium, having been involved with the herbarium since before 1950. As early as 1951 he was collaborating with Santapau on a note published in the The Journal of the Bombay Natural History Society¹² Bole was one of the editors of The Journal of the Bombay Natural History Society from 1971 to 1998.  Bole was still active in field work at the age of 82.⁸

So, I would guess that the first and commonly-grown European clone of Boucerosia frerei is originally from Vajragad Hill Fort via the gardens of Hermenegild Santapau at St Xavier's College in Mumbai. Supplied to Maurice Mason by PV Bole. How soon that was followed by plants sent to Edgar Lamb by Mr Ghoghari of Ahmedabad is still a mystery to me and they still might have been the earlier introduction. 


There is a plant offered for sale from many European growers that is referred to as Frerea indica x Caralluma europaea. A few offered as the hybrid seem to be the pure Frerea indica, misidentified. A few offered as the pure Frerea indica seem to be the hybrid, misidentified. As the Frerea is mentioned first, it was the seed parent and the Caralluma was the pollen donor.

Flowers of the hybrid Frerea indica x Caralluma europaea
Photos kindly supplied by Yael Bar Natan Gayer ©

Jerry Barad commented on this hybrid, as reported by the excellent website. The Marnier collection would be Les Cèdres, see above.¹⁰⁴
A similar hybrid occurred by fly pollination at the Marnier collection many years ago and has been circulating in Europe as Frerea indica x Caralluma europaea. When I visited that garden the curator, Mr. Hebding, advised me that the name was a guess on their part as no controlled artificial pollinations were being done.
Obviously they knew that the Frerea was one parent as it produced the seeds. Which of their Caralluma species was the "father" is uncertain. Though from the look of the flowers Caralluma europaea is a good educated guess.

Habit of the hybrid Frerea indica x Caralluma europaea
Note that the leaves are much smaller than those 
of Frerea but are clearly true leaves.
Photos kindly supplied by Yael Bar Natan Gayer ©

The Alsatian gardener René Hebding had worked at Les Cèdres from 1960 to 2000. Hebding was internationally respected for the depth and breadth of his knowledge about the tens of thousands of species under his care at Les Cèdres. Two plants were named after him: Senecio hebdingii and Dyckia hebdingii. In the 1990s, Hebding was the author of several plant names with Werner Rauh, including Aloe anivoranoensis (Rauh & Hebding) L.E.Newton & G.D.Rowley and Pitcairnia subfuscopetala Rauh & Hebding.

Frerea indica x Caralluma europaea in flower and fruit.
Used by the generous global permission of the late Chris Moore, 
from his incredible gallery⁶¹ Asclepiad Exhibition

Another variety of this hybrid can be seen at Chris Moore's Asclepiad Exhibition, including the photo above with an immature twin fruit. In 2003, the International Asclepiad Society offered seed from Chris Moore of this hybrid on their seedlist.⁶⁵

In the second half of the 1970s Jagdash Singh Sarkaria had given live plants of Frerea indica to the plant collector Gerald "Jerry" S Barad who was based in New York. Barad immediately started hybridising Frerea with various species of Caralluma and Huernia.⁹⁵

Huernia saudi-arabica x Frerea indica has been called x Fernia "Beatrice" after Barad's wife. Barad numbered it 107-83. It was distributed by International Succulent Introductions under the number ISI 92-48.¹⁵ Described as "the stout rich green stems bear glossy, inch-long, elliptical leaves and pretty maroon flowers banded with yellow".¹⁵ A photo can be seen in Barad's article "Pollination of Stapeliads".⁹ 

Frerea indica x Caralluma truncato-coronata was represented by two entries in Sarkaria's list of accessions at his National Cactus, Succulent Botanical Gardens and Research Centre, one of which was given the name "Garnet Stones".⁹⁵ It was distributed by International Succulent Introductions under the number ISI 89-53 (Barad used the number 111-62). Described as "vigorous, slender, apple-green, leafy mat-forming stems and velvety purplish-red flowers".¹⁵ Caralluma truncatocoronata is now known as Boucerosia crenulata

Frerea indica x Caralluma shadhbana var. barhana was represented by three entries in Sarkaria's list of accessions at his National Cactus, Succulent Botanical Gardens and Research Centre.⁹⁵  Caralluma shadhbana var. barhana is now regarded as Monolluma hexagonaA photo can be seen in Barad's article "Pollination of Stapeliads".⁹

 Chris Moore had this plant as a hybrid of unknown parentage, though all 
the other hybrids I have seen in photos have had stunted leaves.
This may be yet another flower form of the true species.
Used by generous global permission of Chris Moore, 
from his incredible gallery⁶¹ Asclepiad Exhibition

Flower structure

A very short video I made to show 
the vibratile hairs waggling.
Worth going full screen.

These vibratile hairs are found on the flowers of some other species of stapeliads. They seem to be associated with stinky fly-pollinated flowers. To me they look like the wings of small flies complete with realistic movement. I presume that this would attract the attention of other flies passing by and looking to lay their eggs on carrion. Perhaps they are reassuring that other flies are there already, not eaten by predators. The Frerea flower is, of course, a lie. There is no prospect of any fly eggs laid on this fake piece of dead meat ever growing to adulthood.

If you want to see some microscopic studies of the surface of the flower and the smell-secreting cells (osmophores) then Plachno et al can satisfy your needs.⁷² They misunderstood the renaming of the plant, referring to it as Boucerosia indica throughout, except for "B. indica (Rowley) Meve & Liede (= Frerea indica Dalz.)". They had the right authors but the wrong name. Boucerosia indica is, of course, a completely different plant also previously called Caralluma indicaBoucerosia hutchinia and Hutchinia indica. The confusion is understandable. The photo and description make it clear that they did mean Boucerosia frerei.

I am not terribly happy with either of these cross-sections 
so I am going to wait until I get another flush of 
flowers before I do a version with a key.

I have posted two other cross-sections of flowers in this family - Stephanotis floribunda and Hoya bella. For an explanation of the pollination strategy of the asclepiads please see the description of that cross-section for Stephanotis floribunda.

The back of one of my flowers.


Caralluma frerei photographed from a cultivated plant 
by the late Chris Moore. The lotus petal shape is quite 
similar to some found in Bhimashankar Wildlife Sanctuary.⁸⁰
Used by kind global permission of Chris Moore, 
from his incredible gallery⁶¹ Asclepiad Exhibition

Some of the variability in flower shape and patterning can be seen in this collection of black and white photos provided by Pundarikakshudu Tetali et al.⁹⁹ The colour photographs at the Flora of Peninsular India Website show an interesting range of patterns and shape in only four plants (the second and third photos are clearly the same flower).⁸⁹ Thirteen colour photographs from the Naoroji Godrej Centre for Plant Research at Shindewadi in Maharashtra show colour variations as well as pattern differences.⁶⁸ One of the photos in Plate VIII in this PhD thesis is also slightly different in the markings.⁴⁰ Another pattern can be seen in the photograph⁹⁸ on page 125 of this PhD thesis, though the photo was taken by the author of the preceding thesis.

The following photos from iNaturalist also show the variability of flower patterns.

Boucerosia frerei on Lenyadri Hill. 
Photograph © Sushant More (2020).⁶³

Boucerosia frerei in Bhimashankar Wildlife Sanctuary. 
Photograph © Mayuresh Kulkarni (2017).⁴²

Boucerosia frerei on Anjaneri Hill, 
the most northerly known population. 
Photograph © Sushant More (2015).⁶²

As far as I know, these plants have not been investigated to see how much genetic variation there is within the species. It would be interesting to know if there are hidden species within the known collections. It is such an easy plant to distinguish from the other stapeliads (from its large leaves and habit) that it might not have been fully appreciated by botanists how much variation has been seen from those fourteen isolated collection sites. There is a tremendous variation in flower patterns just among the plants from Shivneri Hill Fort.

Plant cultivated by Botanical Survey of India.
Photographer Yercaud Elango, 10 October 2017. 
Cropped to plant. From Wikimedia Commons CC BY-SA 4.0

Flower of Boucerosia frerei. Location of plant origin not given.
Photograph: Dr P. Tetali. From Kew Gardens, POWO ⁷³

How the plants got to those hilltops is another open question. It is possible that Frerea was much more widespread and perhaps grew at lower altitudes when conditions were cooler and wetter. The seeds of Frerea have fluff attached and are capable of flight. The hills are very far apart so it is possible that the colonisation of new hills was very rare. 

Vultures have been seen attacking Frerea stems during the dry season. It is thought that they mistake the plant stems for silvery worms. The stems can become detached and probably root where they fall. The author noted⁷⁰ that the possibility that vultures might carry the plant from one hill to another "requires further observation and study".

More than half of the known locations are hill forts and several have ancient Buddhist complexes. It is possible that the plant was deliberately introduced to some hill forts to provide fresh vegetables and for use to cure wounds. Obviously a useful trait for military fortifications. 

Clearly pollination will only occur within plants on the same hillside as the little flies are not going to be flying tens of miles to find another flower.

The flowers above are all from my two plants which I am sure 
are clones of the same original plant. You can see that all 
the flowers have unique sets of five squiggles.
The varying colours are just due to lighting.

This plant obviously made a mistake with this 
flower and only produced 4 petals. It never 
opened fully, remaining concave as it was here.

Though the flying seeds could provide cross-breeding between different hillsides, there is still a significant geographical barrier. Especially as the wild populations do not seem to produce fruit often. Perhaps not as isolated as the tepui of Venezuela, where many species of flowers and animals are found on only one plateau. Of course, some species of plant are very variable in appearance while still being regarded as one species.

There is an apparent gap of 75 km (47 miles) between the southern sites and the northern ones. This may be real or it might be the result of the plant not being noticed on the hills in the middle. It has been remarked that the dormant plant on a vertical cliff face looks like a mass of roots. The plant when dormant or not flowering has been said to be easy to mistake for a stunted Euphorbia neriifolia, which is too common to attract the attention of most botanists.⁹² Perhaps there are also cryptic species among Euphorbia neriifolia

Hopefully, this plant is a lot more common than we might assume from the known collections from the wild. 


¹ Abdulali, H "Obituary: Charles McCann, 1899-1980 (With a plate)" The Journal of the Bombay Natural History Society (1980) 77(3): 494–495     

² Acland, RD signed as RD Bombay "On Frerea indicaThe Journal of the Bombay Natural History Society (1940) 41(3): 679   Accessed 8/7/2020

³ African Succulent Plant Society, The The National Cactus and Succulent Journal Vol. 21, No. 2 (June 1966) Advert Section      Accessed 17/10/2020

⁴ Anonymous "Orchid experts praise Gardens" The Straits Times (1952) 21 March: 10   Accessed 24/10/2020 

⁵ Anonymous "Four judges from abroad" The Singapore Free Press (1952) 28 March, Singapore Flower Show Supplement: 2   Accessed 24/10/2020

⁶ Anonymous "Deceased Fellow: Professor H. Santapau" Indian National Science Academy website  (undated)    Accessed 18/10/2020

⁷ Anonymous "Begonia masoniana" Database of Native Plant in Taiwan website, Academia Sinica (undated)         Accessed 18/10/2020

⁸ Anonymous "Prof. P.V. Bole" Blatter Herbarium website  (undated)        Accessed 31/10/2020

⁹ Barad, GS "Pollination of Stapeliads" Cactus and Succulent Journal (U.S.) (1990) 62(3): 130-140

¹⁰ Bensusan, K et al "On the distribution and habits of Apteranthes joannis (Maire (Plowes)" Asklepios (2009) 104(March): 3-8        I actually owned this one already.

¹¹ Bhosle SV et al "Ethnomedical Knowledge of Plants used by the Tribal people of Purandhar in Maharashtra, India" Ethnobotanical Leaflets (2009) 13: 1353-61     Accessed 21/6/2020

¹² Bole PV & Santapau, H "A note on Neuracanthus sphaerostachyus Dalz." The Journal of the Bombay Natural History Society (1951) 50(2): 428-430    Accessed 16/10/2020

¹³ Bruyns, PV Stapeliads of Southern Africa and Madagascar Volume I (2005, Umdaus Press, Hatfield) Fig. 14        Accessed 2/11/2020

¹⁴ Chandan A "Kailash Puri Obituary" The Guardian (2017) 4th of July      Accessed 28/7/2020

¹⁵ Clark, P "Frerea indica - A Review" Asklepios (1994) 63(December): 23-35 and photos facing page 22   Back issue kindly supplied by the International Asclepiad Society on 30/10/20

¹⁶ Cooke, T "Flora of the Presidency of Bombay" (1903, London: Taylor & Francis) Vol. 1: v & Note     Accessed 9/7/2020

¹⁷ Cooke, T "Flora of the Presidency of Bombay" (1908, London: Taylor & Francis) Vol. 2: 178      Accessed 9/7/2020

¹⁸ Dalzell, NA & Gibson, A "The Bombay Flora: or, Short Descriptions of all the Indigenous Plants hitherto discovered in or near the Bombay Presidency; etc." (1861, Bombay)       Accessed 9/7/2020

¹⁹ Dalzell, NA (collector) & Fitch, WH (artist) with notes added by others "Boucerosia frerei (G.D.Rowley) Meve & Liede" (1863 onwards) Kew Gardens Herbarium sheet      Accessed 27/7/2020

²⁰ Dalzell, NA "Letters" (1863/1864) JSTOR Global Plants Search       Accessed 10/7/2020 (paywall)

²¹ Dalzell, NA "A New Genus of AsclepiadeæThe Journal of the Linnean Society, Botany (1865, London) 8: 10-11   Accessed 18/6/2020

²² Easton, A "Odontoglossum cirrhosum, an interesting and relatively ignored Odontoglossum species." The International Odontoglossum Alliance Newsletter  (2016) 1: 5-9      Accessed 31/7/2020

²³ Elliot, B "Maurice Mason (1912-1993)" The Plantsman (2011, Royal Horticultural Society) New Series 10(1): 58-63    Scan kindly provided by the Membership Office of the Royal Horticultural Society 24/9/2020

²⁴ ENVIS Centre on Floral Diversity, Botanical Survey of India "Frerea indica" (Undated)       Accessed 17/7/2020

²⁵ Fearn, B "Notes on Frerea indicaAsklepios (1994) 63(December): 36-7  Back issue kindly supplied by the International Asclepiad Society on 30/10/20

²⁶ Flowers of India website. Accessed June 20 2020

²⁷ Gibson, A "Report on the state and progress of Agriculture in the Deccan for the year 1842" Journal of the Agricultural & Horticultural Society, India Vol. 2 (1843, Calcutta) 2(6): 286-302     Accessed 26/7/2020

²⁸ Ginns, R "Frerea indica Dalzell" Asclepiadaceae (1974) 1: 1      Back issue kindly scanned by the International Asclepiad Society on 7/12/20

²⁹ Gupta, A "Endangered Plant to be Named ‘Shiv Suman’ to Honour King Shivaji" India's Endangered website (2019)      Accessed 9/7/2020

³⁰ Hardy, DS "Stapeliads: Conditions under which they grow and how they should be cultivated"  Plants & Gardens (1963) 19(3): 15-17    Accessed 30/7/2020
Also published in a special printing as Handbook on Succulent Plants (1963)       Accessed 2/11/2020

³¹ Hooker, JD "The Flora of British India" Vol. IV (1885, L. Reeve & Co., London) Part X (first published June 1883): 76     Accessed 27/7/2020

³² Hunt, R (Revised by Grout, A) "Dalzell, Nicol Alexander" Oxford Dictionary of National Biography (2004)  Accessed online via Manchester Library login 15/9/2020

³³ Irani, NA "N.I. 108" (1954) Herbarium specimen at "Frerea indica Dalz." Flora of Peninsular India Website (2019)       Accessed 28/7/2020

³⁴ Jacobsen, H & Rowley, GD "Some Name Changes in Succulent Plants PART IV" The National Cactus and Succulent Journal (1958) 13(4): 75-78     Accessed 6/7/2020

³⁵ Jadhav, CR "Studies on biodiversity of Pune district: Flowering plants" PhD Thesis, University of Pune (2013) Chapter 9: 472-3     Accessed 30/7/2020

³⁶ Jadhav, SV "Rock-cut cave temples at Junnar : an integrated study" PhD Thesis, Deccan College Post-Graduate and Research Institute, University of Poona (Pune) (1980)     Accessed 26/7/2020

³⁷ Jadhav, R et al "Forest Foods of Northern Western Ghats: Mode of Consumption, Nutrition, and Availability" Asian Agri-History (2015) 19(4): 293–316      Accessed 24/10/2020

³⁸ JSTOR Global Plants "Hardy, David Spencer (1931-1998)" JSTOR Biographies (Last updated 2013)       Accessed 30/7/2020

³⁹ Kamble, MY & Yadav, SR "Asclepiadaceae of Maharashtra" Bulletin of the Botanical Survey of India (2004) 46(1-4): 34-49     Accessed 12/07/2020

⁴⁰ Kamble, MY "Taxonomical studies on Asclepiadaceae and Periplocaceae of Maharashtra"  PhD Thesis, Shivaji University, Kolhapur (2007) Chapter 6: 82-83 & Plate VIII     Accessed 24/10/2020

⁴¹ Kothari, MJ & Moorthy, S "Flora of Raigad district, Maharashtra State" (1993, BSI, Kolkata) 232-233   Not seen.

⁴² Kulkarni, M "Boucerosia frerei" (2018) iNaturalist website

⁴³ Kumbhojkar, MJ et al "Report on a new locality of endemic Frerea indica Dalz. in Satara District" Indian Journal of Forestry (1993) 16: 85-86   Not seen.  

⁴⁴ Lamb, E "Frerea indica. Dalz." Photographic Reference Plate (November 1961) No. 2657  Scan kindly supplied by Chris Leather, curator of Lamb's Photographic Plates, 02/11/2020.

⁴⁵ Lamb, BM et al "Moroccan Asclepiads following autumn storms" Cactus and Succulent Journal (2009) 81(5): 240-255

⁴⁶ Lange, OL & Zuber, M "Frerea indica, a stem succulent CAM plant with deciduous C3 leaves" Oecologia (1977) 31: 67–72     Accessed 9/7/2020

⁴⁷ Lloyd, C "The Expert Gardener: Friends to remember" The Guardian Weekend (2003) 05 July: C67     Accessed 24/10/2020

⁴⁸ Mace, S Personal Communication 27/11/2020

⁴⁹ Marnier-Lapostolle, J "Notes du Jardin Botanique des Cèdres Les Ceropegia (suite) (1)Cactus Revue Trimestrielle (1960) 15(Octobre-Décembre, 68/69): 53-56      Accessed 12/07/2020

⁵⁰ Marshall Woodrow, G "The Flora of Western India. Part I" The Journal of the Bombay Natural History Society (1897) 11: 118-130     Accessed 8/7/2020

⁵¹ Marshall Woodrow, G "The Flora of Western India Part V" The Journal of the Bombay Natural History Society (1898) 12(1): 168    Accessed 20/6/20

⁵² Martin, MJ "Stapeliads" Cactus and Succulent Journal of Great Britain (1966) 28(May, 2): 38-39        Accessed 31/10/20

⁵³ Marshall Woodrow, G "Four interesting Bombay Plants" The Journal of the Bombay Natural History Society (1903) 15: 363-364 + plates     Accessed 8/7/2020

⁵⁴ Mason, LM (collector) "Frerea indica Dalz." (1959) Kew Gardens Herbarium spirit collection      Accessed 1/8/2020

⁵⁵ McCann, C "Additions to the description of Frerea indica Dalz. (Asclepiadaceae) and some observations on the species." The Journal of the Bombay Natural History Society (1939) 41(1): 143-145       Accessed 9/7/2020

⁵⁶ Meve, U & Liede, S  "A molecular phylogeny and generic rearrangement of the stapelioid Ceropegieae (Apocynaceae-Asclepiadoideae)" Plant Systematics and Evolution (2002) 234: 171–209    ResearchGate           Accessed 16/7/2020

⁵⁷ Meyer, FG "Plant Explorations" Crop Research Service, Agricultural Research Service of the United States Department of Agriculture ARS 34-9 (1959) 81-82     Accessed 30/7/2020

⁵⁸ Mishra, DK & Singh, NP "Frerea indica Dalz. (Asclepiadaceae) - a Critically Endangered Plant, now Collected from Ahmednagar District, Maharashtra" Nelumbo (2000) 42(1-4): 157-159            Accessed 12/07/2020

⁵⁹ Mishra DK & Arya KR "Frerea indica Dalz. (Asclepiadaceae): a palaeoendemic plant of Maharashtra. State, India" Geophytology (2010) 38(1-2): 101-104     Researchgate        Accessed 23/6/2020

⁶⁰ Moholkar SM "Floristic Diversity of Some Hills of Mahadev Ranges" PhD Thesis, Shivaji University, Kolhapur (2018) Chapter 4: 484 + Appendix Plate 30     Accessed 14/07/2020

⁶¹ Moore, C "Caralluma f-p (inc. Frerea indica)" Asclepiad Exhibition website, maintained as a tribute to the late Chris Moore. (2009)      Accessed 2/12/2020

⁶² More, S "Boucerosia frerei" (2019) iNaturalist website    Accessed 14/10/20    Also a personal communication on 27/12/2020.

⁶³ More, S "Boucerosia frerei" (2020) iNaturalist website

⁶⁴ Mukerjee, T & Garg, S "A note on Frerea indica Dalz. (Asclepiadaceae)" Journal of Economic and Taxonomic Botany (1984) 5(4): 968        Not seen by me, used as a reference by Clark. This issue is not even indexed on the website of the journal. I am sure it does exist in various libraries.  

⁶⁵ Mulej, I "Seedlists" International Asclepiad Society website (undated)   Accessed 2/12/2020

⁶⁶ Nandikar, MD et al "Floristic enumeration of Torna Fort (Western Ghats, India): a storehouse of endemic plants" Journal of Threatened Taxa (2018) 10(7): 11895-11915      Accessed 23/6/2020

⁶⁷ Nandikar, MD et al "Taxonomy and typification of Kalanchoe olivacea and K. bhidei (Crassulaceae)" Rheedea (2019) 29(3): 197-208    Accessed 9/12/2020

⁶⁸ Naoroji Godrej Centre for Plant Research "Frerea indica Dalz." Haritarium (undated, last reference 2004)      Accessed 28/7/2020

⁶⁹ Naseem, I "10 years on, Shindel Makudi is no longer an endangered plant: study" The Indian Express (2010) June 29     Accessed 6/7/2020

⁷⁰ Nayar, MP "Endemic Flora of Peninsular India and its Significance" The Bulletin of the Botanical Survey of India (1980) 22(1-4): 12-23      Accessed 6/12/2020

⁷¹ Padhye, A et al "Butterflies of Northern Western Ghats: A Compilation of Checklists" ELA Journal (2013) 2(1): 3-22      Accessed 28/7/2020

⁷²  Plachno, BJ et al "Can a stench be beautiful? – Osmophores in stem-succulent stapeliads (Apocynaceae-Asclepiadoideae-Ceropegieae-Stapeliinae)" Flora (2010) 205: 101-105      Accessed 12/07/2020

⁷³ POWO (2019) Plants of the World Online. Facilitated by the Royal Botanic Gardens, Kew. Published on the Internet
Frerea    Accessed 20/6/2020
Abutilon      Accessed 9/7/2020

⁷⁴ Prain, D "A New Curcuma from the Deccan" The Journal of the Bombay Natural History Society (1898) 11(3): 463-464 + plate   Accessed 9/7/2020

⁷⁵ Punekar, S quoted by Nitnaware, H in "Endangered flower to get vernacular name" Pune Mirror (2019) May 23    Accessed 21/6/2020

⁷⁶ Puri, KK "A Brief Autobiography" Sikh Heritage Website (Undated)       Accessed 28/7/2020     

⁷⁷ Putnam, EW (as E.W.P.) "Ron Ginns F.N.C.S.S. 1896-1976" The National Cactus and Succulent Journal (1977) 32(1, March): 2-3   Accessed 8/12/2020

⁷⁸ Raghavan, RS "A Short Note on Frerea indica Dalz." Current Science (1976) 45(1): 36  Letter to journal dated September 10, 1975.     Accessed 6/7/2020 (Registration free)

⁷⁹ Rahangdale, SR & Rahangdale, SS "Potential Wild Edible Plant Resources from Maharashtra: Future prospects for their conservation and improvement" Life Sciences Leaflets (2014) 57: 84       Accessed 25/10/2020

⁸⁰ Rahangdale, SS & Rahangdale, SR "Floristic diversity of Bhimashankar Wildlife Sanctuary, northern Western Ghats, Maharashtra, India" Journal of Threatened Taxa (2017) 9(8): 10493-10527      Accessed 26/7/2020

⁸¹ Rauh, W "Frerea indica Dalz., eine wenig bekannte Stapeliee aus IndienKakteen und andere Sukkulenten (1961) 12(2): 24-25     Accessed 12/07/2020

⁸² Reiß, J "Caralluma frerei ROWL. (Frerea indica DALZ.), ein bemerkenswerter Vertreter der sukkulenten Asclepiadaceen" Kakteen und andere Sukkulenten (1967) 18(5): 90-92     Accessed 12/07/2020

⁸³ Rose, H "Floraisons les plus intéressantes observées dans les serres du Muséum pendant les anneés 1964 et 1965" Bulletin du Muséum National de Histoire naturelle Série 2 (1966) 38(2): 208-216     Accessed 12/07/2020

⁸⁴ Rowley, GD "Whitsun Safari" The National Cactus and Succulent Journal (1958) 13(3): 47-52      Accessed 6/7/2020

⁸⁵ Rowley, G.D. "Frerea: genus or section?" Asclepiadaceae (1974) 3: 2-3     Back issue kindly scanned by the International Asclepiad Society on 26/11/20

⁸⁶ Rowley, GD "...the following comment" Asklepios (1994) 63(December): 37 and photos on page 25 of Phil Clark's article in the same issue. Back issue kindly supplied by the International Asclepiad Society on 30/10/20

⁸⁷ Rowley, GD "A History of Succulent Plants" (1997, Strawberry Press, California) p. 375     Scan kindly provided by Kew Gardens Library on 22/10/2020

⁸⁸ Royal Botanic Gardens Edinburgh Website "The Dapuri Drawings" (2002?)    Accessed 26/7/2020

⁸⁹ Sankara Rao, K et al "Frerea indica Dalz." Flora of Peninsular India Website (2019)       Accessed 28/7/2020

⁹⁰ Santapau, H "New Record for Frerea indica Dalz. in Bombay Province." The Journal of the Bombay Natural History Society (1950) 49(2): 801-2     Accessed 8/7/2020

⁹¹ Santapau, H "The Flora of Purandhar: or, An enumeration of all the phanerogamic plants discovered in Purandhar during the years 1944-1956" (1957?, Oxford Book and Stationery Co., New Delhi) 81   Scan kindly provided by Kew Gardens Library on 16/10/2020

⁹² Santapau, H & Irani, NA "The Asclepiadaceae & Periplocaceae of Bombay" University of Bombay Botanical Memoirs, No. 4(1960 or 1962?, University of Bombay) 44-45   Scan kindly provided by Kew Gardens Library on 10/9/2020

⁹³ Sarkaria, JS "In search of Frerea indica Dalz." Asclepiadaceae (1980) 21: 29-34     Back issue kindly scanned by the International Asclepiad Society on 7/12/20

⁹⁴ Sarkaria, JS “Indian Succulent Log—Parts V, VIII, IX” Journal of National Cactus & Succulent Society of India (1983) III: 76-82, (1985) V: 66-79, (1986) VI: 44-64      Accessed 27/7/2020

⁹⁵ Sarkaria, JS “Stapeleiae Collection Record up to July 15th 1991" National Cactus, Succulent Botanical Gardens and Research Centre: Sector 5, Panchkula, Haryana, India (1991) 13      Accessed 27/7/2020

⁹⁶ Sawant-Kulkarni, N & Shinde, V "Study of the Military Architectures and Remains of the Shivneri Fort, Maharashtra, India" Bulletin of the Deccan College Post-Graduate and Research Institute (2015) 75: 125-134     Accessed 26/7/2020

⁹⁷ Stillwell, M "Mrs Stillwell on Succulents" Cactus and Succulent Journal of Great Britain (1963) 25(1): 20-22     Accessed 24/10/2020 

⁹⁸ Surveswaran, S "Molecular phylogenetics and medicinal plants of Asclepiadoideae from India" PhD Thesis, University of Hong Kong (2008) 124-127      Accessed 6/12/2020

⁹⁹ Tetali, P et al "Studies in the status and conservation of Frerea indica Dalz" Journal of the Bombay Natural History Society (1997) 94(1): 115-121   Accessed 22/3/2020

¹⁰⁰ Tetali, P et al "Association of Frerea indica Dalz., an endangered plant species with Euphorbia neriifolia L. and its importance in habitat conservation" Current Science (1997) 73(7): 563-565     Accessed 22/3/2020

¹⁰¹ Tyrrell, CR (Lush) "Thoughts and Observations" The National Cactus and Succulent Journal (1968) 23(2, June): 51    Accessed 8/12/2020

¹⁰² Tyrrell, C.R. "Further notes on Frerea indica" Asclepiadaceae (1974) 3: 11      Back issue kindly scanned by the International Asclepiad Society on 26/11/20

¹⁰³ Vasant, S "Yavanas in Western India" Bulletin of the Deccan College Post-Graduate and Research Institute (1988-89) 47/48: 331-338       Accessed 26/7/2020

¹⁰⁴ Viljoen, L & D'Alessi, F ""  Accessed since 2011

¹⁰⁵ Waman, MB "Ecologial studies of Kalsubai, Harishchandragarh, Ratangarh forest" PhD Thesis, Savitribai Phule Pune University (2005) Chapter 5: 101 + Plate 17       Accessed 14/07/2020

¹⁰⁶ Waman, MB & Khyade, MS "Ethnobotanical Uses of Some Plants of Families Apocynaceae and Asclepiadaceae from the Northwestern Region of Ahmednagar" Plant and Human Health, Volume 1: Ethnobotany and Physiology Eds. Ozturk, M & Hakeem, KR (2018) p. 575    Accessed 24/10/2020

¹⁰⁷ Watve, A "Maharashtra" Rocky Plateau site profiles,  Rocky Plateaus Networking Project,
Biome Conservation Foundation (2013)       Accessed 28/7/2020

¹⁰⁸ Woodward, A. "Caralluma frerei (Dalz.) Rowley" Asclepiadaceae (1974) 1: 5-6      Back issue kindly scanned by the International Asclepiad Society on 26/11/20