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|Vol. 10(8), pp. 8-15||The McAllen International Orchid Society Journal||August 2009|
"Few flowers are more remarkable in their structure, or contain more curious appendages than this." Conrad Loddiges, 1818
Botany is a long conversation, one taking place over centuries. Sometimes one participant in a conversation about a specific plant will be talking to another who is long dead. Such are the ways of science.
Bonatea speciosa, our subject here, has been scientifically conversed over since the end of the 18th century. It must have, of course, been seen by man long before that, as it is such a noticeable plant when in flower. The First People who encountered it were the San (or Bushmen), hunter-gatherers who inhabited what is now South Africa for more than 150,000 years. They are now long extinct in the distribution range of B. speciosa and we know nothing of their associations with this plant. The Khoekhoen (Khoi-khoi or Hottentots), nomadic herdsmen who appeared some 2,000 years ago, were inhabitants of the southern and western Cape 350 years ago when the first European settlement of South Africa commenced. Pioneers unfortunately tend to have neither the time nor the inclination to be students - the more basic needs of life being uppermost in their minds - and thus a great opportunity to learn was lost before the last culturally-adherent Khoi-khoi were hounded into oblivion through intermarriage, interracial aggression, disease and religious propaganda. Their blood still runs thick in the veins of South Africa, but their culture is now virtually unknown.
The Bantu-speaking tribes from central Africa migrated within the last thousand or so years into South Africa; with firstly the Sotho group moving down the uplands of the eastern centre of the country, and then from about 700 years ago, the Nguni group moved down the east coast. Some San and Khoi-khoi were absorbed into the migrating Bantu-speaking tribes, notably the medicine men. The long-gathered knowledge of these latter professionals was co-opted into the still-persisting culture of especially the Nguni tribes, and remains today. Bonatea speciosa does not appear to have any current medicinal uses amongst the Nguni, and thus we can infer that it was probably not considered of any use by the San or Khoi-khoi. None of the above kept any form of record for future reference aside from oral tradition.
And thus we come to records - the very means by which the long conversation is being conducted. The first record of this plant, as Orchis speciosa, is to be found in the work of the younger Linnaeus (L.f.); Supplementum Plantarum of 1782.
This Latin is fairly easy to translate; just bear in mind that some of those letters that look like f's are long s's - hence scapo folioso (leafy scape), etc. It is right here that we find a misinterpretation of the structure of the flower - 'labello quinquepartito' (a five-part lip). Now is thus a good time to look at the structure of the complicated flower of Bonatea speciosa; every South African learner orchid judge's nightmare.
Fig. 1. B. speciosa flower with parts labeled. Digital jpg photo from author, Sun-29Jul09.
Fig. 2. Five-whorl petloid monocot diagram. Digital jpg photo from author, Sun-29Jul09.
It can be seen that what was initially interpreted as a five-part lip is actually a three-part lip, fused basally with the anterior lobes of the bizarre divided petals, one on each side. And even if the status of the petals was understood, then there was the matter of deciding exactly what those stigmatic arms represented. So you can't really blame L.f., with all the fusion between the various floral parts, this is a particularly difficult flower to get one's mind around (Fig. 1) (Fig. 2).
As can be seen, the specimen described by L.f. was one collected by the Swede Carl Peter Thunberg. Thunberg was a student of the elder Linnaeus, one who was to become a 'Linnaean Apostle'. Following his graduation from Uppsala University, Thunberg wanted desparately to go and explore the botany of Japan, but he encountered the problem that only Dutch nationals were allowed on the Emperor's soil. In order to pass as a Dutchman, he came and spent a few years in the Cape of Good Hope, where Dutch was the lingua franca (Afrikaans, a language primarily derivative of 18th century Dutch, is still very widely spoken here in South Africa and is my second language). During his time here, he travelled and collected extensively until he was sufficiently Dutchified to proceed to Japan and botanise there while serving as a medical officer. He spent a year in Japan and some time in Java and Sri Lanka and his travels lasted in all some nine years. Thunberg unfortunately never annotated his specimens with locality data, aside for the annotation 'C.B.S.' (Caput Bonae Spei = Cape of Good Hope), and thus we do not have an actual type locality for 'Orchis speciosa'. On his return to Sweden, Thunberg commenced writing up his botanical acquisitions, and listed this species in his 1794 Prodromus thus: "Orchis speciosa. Orchis labello 3-partito: laciniis flexuosis, foliis ovatis". So it would appear that he had indeed seen the light with regard to the basic architecture of the lip (more about this later).
Fig. 3. The first published illustration of Bonatea speciosa, that from Jacquin's Plantarum rariorum horti caesari schoenbrunnensis of 1804. jpg photo from author, Sun-29Jul09.
In 1804, Jacquin published the first illustration of this species in his Plantarum rariorum horti caesari schoenbrunnensis, (Fig. 3) based on a specimen growing in a greenhouse in the Royal Garden at the Palace of Schönbrunn, near Vienna. And what a magnificent plate this is, rendered via copper plate intaglio by one of his two artists, Johannes Scharf or Martin Sedelmayer. In the all-Latin text, Jacquin pointed out that dried specimens from the Cape had longer scapes and were twice as floriferous as the cultivated one he had depicted. His long and complicated Latin description written in what I call 'Jacquinese' - a language conceptually difficult to decypher - does describe the lip as 'quinquipartitum'.
In 1805, Carl Ludwig Willdenow had volume 4 of his reworking of Linnaeus's Species Plantarum published, in which he erected a new genus for this plant. He named it in honour of the then late Professor Giuseppe Antonio Bonato of Padua in Italy - a man important as the Prefect of the Botanic Garden in Padua and whose personal herbarium, together with that of his predecessor Giovanni Marsili, formed the basis of the Padua Herbarium (PAD), now housing in excess of a half million specimens. Unfortunately this choice was poor - Bonato never had anything to do with this plant or South African botany in general. Once again, the labellum was given as quinquipartitum. Willdenow also erected the related genus, Habenaria, in the same publication.
Thunberg's Flora Capensis of 1807, in which we find an extensive description of this species in Latin, surprisingly still has this plant labelled Orchis speciosa and the lip is described as '5-partum' - so perhaps the earlier Thunberg description of a 3-part lip was a printer's error and not a flash of brilliance. At least here we find a description of where the plant was observed by Thunberg, namely (tr.) "near several rivers, (such) as Kafferskuils River (now renamed the Goukou River), Krom River, Seekoei River & in Mossel Bay." The information on Bonatea speciosa is unchanged through the various editions of Thunberg's work; I looked it up in the 1807 original, 1818 Editio nova and in the 1823 Schultes edition.
Fig. 4. Map of the 1772 and 1773 Thunberg journeys to what is now the Eastern Cape.
Below is a map (Fig. 4) showing the routes of the two journeys undertaken by Carl Peter Thunberg to what is now the Eastern Cape in 1772 and 1773. ++++ indicates the first journey when he was accompanied by the master gardener, Johann Andreas Auge and oooo the second on which he was accompanied by the English plant collector, Francis Masson. 1 is the Goukou (formerly Kafferskuil) River - Thunberg probably saw B. speciosa in the area now occupied by the town called Riversdale; 2 is Mossel Bay; 3 is the Krom River and 4 is the Seekoei River.
In 1818, a plate of this plant and a brief note in English was published as t. 284 in volume 3 of the Botanical Cabinet of Conrad Loddiges & Sons. Aside from mentioning the nomenclature of this plant and something about the culture in their greenhouse, Loddiges said "Few flowers are more remarkable in their structure, or contain more curious appendages than this. It is truly an astonishing production, deserving the minutest examination ; and irresistibly commanding our utmost admiration of the skill displayed in its formation by the Infinite Creator." The plate of the spike is well-executed and life-like, but lacks the crispness of the Jacquin plate.
Sometime around 1820, cultivation of this species commenced at Kew - unfortunately too late for inclusion in the Hortus Kewensis (1810-1813) of William Townsend Aiton, the then superintendant of the Royal Gardens, which included Kew. Terrestrial orchids are usually difficult to cultivate; however Bonatea speciosa is one of the exceptions, growing under a wide variety of greenhouse conditions in any well-drained sandy compost; with some regard being paid to its seasonality.
Fig. 5. The original 1826 Greville plate from Curtis's Botanical Magazine (t. 2926). (jpg photo from author, Sun-29Jul09)
In 1826, a plant was forwarded from Kew to the botanical gardens in Edinburgh, Scotland, and upon flowering, this plant was described for Curtis's Botanical Magazine (t. 2926) by Prof. Robert Graham, the Scots-born Professor of Botany at the University of Edinburgh and Regius Keeper of the RBG, Edinburgh. The accompanying plate was executed by Dr. Robert Greville, another Scots botanist (Fig. 5).
The long and comprehensive description of the plant in English (all earlier description were rendered in Latin) is unfortunately conceptually erroneous in part (including, as you guessed, "Lower segment (labellum) fleshy, unequally divided into five; ....") and although the drawing is botanically most useful, it lacks the great artistic beauty of the Jacquin plate.
In 1831, Robert Brown, the most intellectual botanist of that era, published some of his findings on fertilisation in orchids and asclepiads. He was the man who finally clarified the structure of the flower of Bonatea speciosa:- "It would seem that in Bonatea the extraordinary development and complete separation of these lateral stigmata, have effectually concealed their true nature; and accordingly they have uniformly been considered as forming parts or appendages of the labellum, with which indeed their bases cohere. That they are really stigmata, however, I have proved by a careful examination of the tissues of their secreting surface, by the action of the pollen artificially applied to this tissue, by the descent of its tubes, hereafter to be described, along the upper surface of the styles which is destitute of epidermis, and by the consequent enlargement of the ovarium."
The amazing care and thoroughness taken by Brown in proving the nature of these stigmatic arms is indicative of the great quality of his work, as well as the thought he put into everything he did. Much of the pioneering work he did, particularly the microscopy, is extraordinary, as he did not have the shoulders of his forerunners upon which to stand. Luckily he scarcely needed them. It was in this same paper that Brown first named and described the cell nucleus.
Brown did not always get everything right. Regarding plants of the genus Ophrys, which at least in Ophrys apifera are often apomictic, he said: "Hence it may be conjectured, that the remarkable forms of the flowers in this genus are intended to deter not attract insects, whose assistance seems to be unnecessary......."
It took until 1877 for someone to say, in regard to Ophrys, "To me ... it seems obvious that the male insect came to the flower mistaking it for a female". This, the first recognition of pseudocopulation as a mechanism to effect pollination, was written by H.G. Wells, later to become the author of The War of the Worlds, and other works of science fiction. The right sort of mind was obviously needed to see this.
After that small diversion, let us get back to Bonatea. In the paper delivered in 1832, Brown spent considerable space discussing his 'mucous cords', which he knew had something to do with fertilisation, but which he considered different to pollen tubes. In this, he appears to have been wrong - they were ultimately both pollen tubes in various states of development - but then again, he had not the shoulders of predecessors..... Most of this work had been done with Bonatea speciosa as subject.
Between 1830 and 1840, the great John Lindley, then Professor of Botany at the University College, London, published his The Genera and Species of Orchidaceous Plants. Other workers had made suggestion as to how the various orchid genera related to one another, but Lindley was the first to classify the orchids systematically, coming up with an arrangement that covered the entire family. In his 4th tribe, the Ophrydeae, in a section of the above work published in October 1835, he placed the genus Bonatea, including Bonatea speciosa amongst a whole range of other plants from Brazil, India and Madagascar which no longer find a place in this genus. Lindley's Ophrydeae - now known as the Orchidoideae - include a majority of the terrestrial orchids from all continents. In his key at the beginning of the Ophrydeae section, he classified Bonatea close to Habenaria, Ate, Diplomeris and Cynorchis. Modern classifications place Bonatea near some of these same genera today.
Fig. 6. The 1849 plate from A Century of Orchidaceous Plants, 'attributed' to W. Fitch. (jpg photo from author, Sun-29Jul09)
Published in 1849, plate 95 in W.J. Hooker's A Century of Orchidaceous Plants depicted Bonatea speciosa. This plate was identical in almost every respect to that published 23 years before in Curtis's Botanical Magazine, except that less of the plate was hand-coloured (Fig. 6).
A careful examination will also reveal that this plate was lithographed, rather than being the product of copper plate intaglio, but the most interesting feature appears on the signature line at the bottom, where we are informed that the image was drawn (delineatus]) by W. Fitch. There was no need to draw this image, that had already been done. Perhaps Fitch lithographed the image, he was famous as an artist/lithographer and his life's output would top 10,000 botanical plates. There is, however, no mention on the plate of the original artist, Greville. In those days, copyright was not yet a legal concept, and much copying of illustrative material took place, however it is hard to imagine a more blatant case of plagiarism, and what's more one practiced by the Director of the RBG, Kew.
Thus by the middle of the 19th century, the information on Bonatea speciosa included descriptions; material on distribution, nomenclature and classification; some idea of its anatomy; mechanisms involving fertilisation; its culture; and its beauty was demonstrated by various botanical illustrations.
In the next part we will look at later work into a deeper understanding of this species.
Grateful thanks to Bill Liltved for kindly reviewing the script and to Missouri Botanical Garden http://www.botanicus.org for the majority of the images, so generously offered by them for all to use.
Brown, R. 1831. On the Organs and Mode of Fecundation In Orchideae and Asclepiadeae. Trans. Linn. Soc. London 16 (1833): 685-738.
Brown, R. 1832. Additional Observations on the Mode of Fecundation in Orchideae. Trans. Linn. Soc. London 16 (1833): 739-745.
Graham, R. 1829. Bonatea speciosa. Showy Bonatea. Curtis's Botanical Magazine 3 (new series): t. 2926.
Hooker W.J. 1849. A Century of Orchidaceous Plants. London: Reeve, Benham, and Reeve. t. 95.
Jacquin, N. J. 1804. Plantarum rariorum caesari schoenbrunnensis. Viennae: C. F. Wappler. 4: t. 451.
Lindley, J. 1834. The Genera and Species of Orchidaceous Plants. London: Ridgways, Piccadilly. p. 327.
Linné, Carl von (L.f). 1782 ('1781') Supplementum plantarum Systematis vegetabilium. Brunsvigae: Impensis Orphanotrophei. p. 401.
Loddiges, C. 1818. Bonatea speciosa. Loddiges's Botanical Cabinet 3: t. 284.
Thunberg, C. P. 1794. Prodromus plantarum Capensium :quas in promontorio Bonæ Spei Africes. Upsaliæ: J. Edman. 1: 4
Thunberg, C.P. 1807. Flora Capensis. Upsaliae: J. F. Edman. 1: 43
Thunberg, C.P. 1818. Flora Capensis, Editio nova. Hafniae: G. Bonnierum. 1: 43
Thunberg, C.P. 1823. Flora Capensis, (Schultes edition). Stuttgardtiae: J.G. Cottae. p. 5
Willdenow, K. L. 1805. Caroli a Linné Species plantarum. Berolini : G.C. Nauk, 4(1): 43