On the natural order of plants called Proteaceae
Encyclopedia
On the natural order of plants called Proteaceae, also published as "On the Proteaceae of Jussieu", was a paper written by Robert Brown
Robert Brown (botanist)
Robert Brown was a Scottish botanist and palaeobotanist who made important contributions to botany largely through his pioneering use of the microscope...

 on the taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

 of the plant family Proteaceae
Proteaceae
Proteaceae is a family of flowering plants distributed in the Southern Hemisphere. The family comprises about 80 genera with about 1600 species. Together with the Platanaceae and Nelumbonaceae they make up the order Proteales. Well known genera include Protea, Banksia, Embothrium, Grevillea,...

. It was read to the Linnean Society of London
Linnean Society of London
The Linnean Society of London is the world's premier society for the study and dissemination of taxonomy and natural history. It publishes a zoological journal, as well as botanical and biological journals...

 in the first quarter of 1809, and published in March 1810. It is significant for its contribution to the systematics
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...

 of Proteaceae, and to the floristics
Floristics
Floristics is a subdomain of botany and biogeography that studies distribution and relationships of plant species over geographic areas.The term is not to be confused with floristry....

 of Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, and also for its application of palynology
Palynology
Palynology is the science that studies contemporary and fossil palynomorphs, including pollen, spores, orbicules, dinoflagellate cysts, acritarchs, chitinozoans and scolecodonts, together with particulate organic matter and kerogen found in sedimentary rocks and sediments...

 to systematics.

Background

Brown had been botanist during Matthew Flinders
Matthew Flinders
Captain Matthew Flinders RN was one of the most successful navigators and cartographers of his age. In a career that spanned just over twenty years, he sailed with Captain William Bligh, circumnavigated Australia and encouraged the use of that name for the continent, which had previously been...

' circumnavigation of Australia, and since returning in England
England
England is a country that is part of the United Kingdom. It shares land borders with Scotland to the north and Wales to the west; the Irish Sea is to the north west, the Celtic Sea to the south west, with the North Sea to the east and the English Channel to the south separating it from continental...

 in 1805 he had been preparing descriptions of the specimens collected during the voyage. Brown's intention was to publish a flora
Flora
Flora is the plant life occurring in a particular region or time, generally the naturally occurring or indigenous—native plant life. The corresponding term for animals is fauna.-Etymology:...

 of Australia, but this was still incomplete in September 1808 when Jonas Dryander asked Brown to write a monograph on the Proteaceae so that Dryander could use Brown's names in a new edition of Hortus Kewensis
Hortus Kewensis
Hortus Kewensis, or a Catalogue of the Plants Cultivated in the Royal Botanic Garden at Kew by William Aiton was a 1789 catalogue of all the plant species then in cultivation at the Royal Botanic Gardens, Kew, which constituted the vast majority of plant species in cultivation in all of England...

. Brown immediately set to work researching the topic.

In preparing the paper, Brown had unfettered access to the herbarium of his patron, Sir Joseph Banks
Joseph Banks
Sir Joseph Banks, 1st Baronet, GCB, PRS was an English naturalist, botanist and patron of the natural sciences. He took part in Captain James Cook's first great voyage . Banks is credited with the introduction to the Western world of eucalyptus, acacia, mimosa and the genus named after him,...

, which included the specimens he had collected during Flinders' voyage, and which contained the most extensive collection of Proteaceae in the world. He was also given access to James Edward Smith
James Edward Smith
Sir James Edward Smith was an English botanist and founder of the Linnean Society.Smith was born in Norwich in 1759, the son of a wealthy wool merchant. He displayed a precocious interest in the natural world...

's Linnean collection; the herbarium of Aylmer Bourke Lambert
Aylmer Bourke Lambert
Aylmer Bourke Lambert was a British botanist, one of the first fellows of the Linnean Society.He is best known for his work A description of the genus Pinus, issued in several parts 1803-1824, a sumptuously illustrated folio volume detailing all of the conifers then known...

; the living collections of George Hibbert
George Hibbert
George Hibbert was an eminent English merchant, politician, slave- and ship-owner, amateur botanist and book collector. With Robert Milligan, he was also one of the principals of the West India Dock Company which instigated the construction of the West India Docks on London's Isle of Dogs in 1800...

; the herbarium of William Aiton
William Aiton
William Aiton was a Scottish botanist.Aiton was born near Hamilton. Having been regularly trained to the profession of a gardener, he travelled to London in 1754, and became assistant to Philip Miller, then superintendent of the Chelsea Physic Garden...

, which contained the collections of Francis Masson
Francis Masson
Francis Masson was a Scottish botanist and gardener, and Kew Gardens’ first plant hunter.Masson was born in Aberdeen. In the 1760s he went to work at Kew Gardens as an under-gardener. Masson was the first plant collector to be sent from Kew by the newly-appointed director Sir Joseph Banks...

; and the collections of William Roxburgh
William Roxburgh
William Roxburgh was a Scottish surgeon and botanist. He has been called the Father of Indian Botany.-Early life:Roxburgh was born at Underwood in the parish of Craigie, Ayrshire. He studied medicine in Edinburgh...

. In addition he was accorded a brief examination of the specimens collected by Jacques Labillardière
Jacques Labillardière
Jacques-Julien Houtou de Labillardière was a French naturalist noted for his descriptions of the flora of Australia. Labillardière was a member of a voyage in search of the La Pérouse expedition...

 at Esperance Bay
Esperance Bay
Esperance Bay is a bay on the south coast of Western Australia. Nominally located at , it is the site of the town of Esperance.The bay was discovered on 9 December 1792 by a French expedition under Bruni d'Entrecasteaux, which sailed in search of the lost expedition of Jean-François de Galaup,...

, on the southwest coast of Australia, where Brown had not collected.

Content

Brown's paper was read to the Linnean Society of London in four parts, on 17 January, 7 February, 21 February and 7 March 1809. The Council of the Society approved it for publication on 2 May, but it did not appear in print until 8 March 1810. It was published as a separate offprint under the title On the natural order of plants called Proteaceae, and then appeared in Volume 10, Part 1 of Transactions of the Linnean Society of London, under the title On the Proteaceae of Jussieu. It was illustrated by two Ferdinand Bauer
Ferdinand Bauer
Ferdinand Lucas Bauer was an Austrian botanical illustrator who travelled on Matthew Flinders' expedition to Australia.-Biography:...

 plates, depicting Knightia excelsa and Dryandra formosa (now Banksia formosa
Banksia formosa
Banksia formosa, commonly known as Showy Dryandra, is a shrub endemic to Western Australia. It was known as Dryandra formosa until 2007, when all Dryandra species were transferred to Banksia by Austin Mast and Kevin Thiele....

).

The paper begins with a digression, presenting Brown's observations on undeveloped Asclepiadaceae
Asclepiadaceae
According to APG II, the Asclepiadaceae is a former plant family now treated as a subfamily in the Apocynaceae...

 flowers as evidence of the importance of studying developing flowers as well as mature ones. It then discusses the biogeography
Biogeography
Biogeography is the study of the distribution of species , organisms, and ecosystems in space and through geological time. Organisms and biological communities vary in a highly regular fashion along geographic gradients of latitude, elevation, isolation and habitat area...

 of Proteaceae, noting the southern hemisphere
Southern Hemisphere
The Southern Hemisphere is the part of Earth that lies south of the equator. The word hemisphere literally means 'half ball' or "half sphere"...

 (that is, Gondwana
Gondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...

n) distribution of the family, and that "[t]he most numerous genera are also the most widely diffused." Brown then discusses the morphology
Morphology (biology)
In biology, morphology is a branch of bioscience dealing with the study of the form and structure of organisms and their specific structural features....

 of the family, including a discussion of pollen
Pollen
Pollen is a fine to coarse powder containing the microgametophytes of seed plants, which produce the male gametes . Pollen grains have a hard coat that protects the sperm cells during the process of their movement from the stamens to the pistil of flowering plants or from the male cone to the...

 shape; according to David Mabberley
David Mabberley
Professor Dr. David John Mabberley, , is a botanist, educator and writer. Among his varied interests is the taxonomy of tropical plants, especially trees of the families Labiatae, Meliaceae and Rutaceae. He is perhaps best known for his plant dictionary The plant-book. A portable dictionary of the...

 this represents "the first major study of the subject, palynology
Palynology
Palynology is the science that studies contemporary and fossil palynomorphs, including pollen, spores, orbicules, dinoflagellate cysts, acritarchs, chitinozoans and scolecodonts, together with particulate organic matter and kerogen found in sedimentary rocks and sediments...

".

The systematics
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...

 section follows, and is most important for the large number of new taxa published therein. 404 species in 38 genera are listed. 18 of the genera are new, and nearly all of these are still upheld. The arrangement of these genera largely follows that introduced by Richard Salisbury in his 1806 The Paradisus Londonensis, although Brown asserts that he arrived at his arrangement independently. Brown does introduce one important concept, however: his division of the Proteaceae into two subfamilies based on whether or not the fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

 dehisces
Dehiscence (botany)
Dehiscence is the opening, at maturity, in a pre-defined way, of a plant structure, such as a fruit, anther, or sporangium, to release its contents. Sometimes this involves the complete detachment of a part. Structures that open in this way are said to be dehiscent...

. This is still considered the fundamental division in the family, though opinion has varied on whether the defining character should be fruit dehiscence or paired flowers.

Overall, the paper was very well received; according to Mabberley, "[t]he paper was masterful and commanded respect for Brown as a brilliant botanist."

Controversy

Shortly after the reading of "On the natural order of plants called Proteaceae", but before it appeared in print, there appeared a publication that started one of the most bitter disputes in 19th century botany. This was On the cultivation of the plants belonging to the natural order of Proteeae
On the cultivation of the plants belonging to the natural order of Proteeae
On the cultivation of the plants belonging to the natural order of Proteeae is an 1809 paper on the Proteaceae family of flowering plants. Although nominally written by Joseph Knight as a paper on cultivation techniques, all but 13 pages consists of an unattributed taxonomic revision now known to...

, ostensibly by Joseph Knight. Despite the title, this paper contained only 13 pages related to cultivation techniques, but over 100 pages of taxonomic revision. Although not explicitly attributed, it was widely believed, and is still so, that this revision was contributed by Richard Salisbury, who had been present at all four of the meetings at which Brown read his paper. The revision contains many of the plant names that Brown had presented to the Linnean Society; for example, the genera Petrophile
Petrophile
Petrophile is a genus of evergreen shrubs, in the protea family Proteaceae, which are endemic to Australia. Commonly known as Conebushes, they typically have prickly, divided foliage and produce prominently-displayed pink, yellow or cream flowers followed by grey, conical fruits...

, Isopogon
Isopogon
Isopogon is a genus of 35 species of mainly low-growing and prostrate perennial shrubs in the family Proteaceae endemic to Australia. They are found throughout Australia, though Western Australia has the greatest variety with 27 of the 35 species found there...

and Grevillea
Grevillea
Grevillea is a diverse genus of about 360 species of evergreen flowering plants in the protea family Proteaceae, native to Australia, New Guinea, New Caledonia, and Sulawesi. It was named in honour of Charles Francis Greville. The species range from prostrate shrubs less than 0.5 m tall to trees...

. Thus Salisbury beat Brown to print, claiming priority for the names that Brown had authored. Salisbury is also accused of having appropriated some of Brown's observations; for example, in the Linnean Society copy of Knight's paper, where Salisbury says that he suspects that the fruit of Persoonia
Persoonia
Persoonia is a genus of 98 species of shrubs and small trees in the tribe Persoonioideae in the large and diverse plant family Proteaceae. In the eastern states of Australia, they are commonly known as Geebungs, while in Western Australia and South Australia they go by the common name Snottygobbles...

is unusual, Brown has pencilled in "He suspects it because he listened very attentively to my paper when read at the Linnean Society."

Salisbury was accused of plagiarism
Plagiarism
Plagiarism is defined in dictionaries as the "wrongful appropriation," "close imitation," or "purloining and publication" of another author's "language, thoughts, ideas, or expressions," and the representation of them as one's own original work, but the notion remains problematic with nebulous...

 and ostracised from botanical circles. Contemporary notes and letters indicate widespread condemnation of Salisbury's actions. For example, Samuel Goodenough
Samuel Goodenough
Samuel Goodenough was the Bishop of Carlisle from 1808 until his death in 1827, and an amateur botanist and collector. He is honoured in the scientific names of the plant genus Goodenia and the Red-capped Robin .-Life:Born at Kimpton, near Weyhill, Hampshire, on 29 April 1743 , he was the third...

 wrote "How shocked was I to see Salisbury's surreptitious anticipation of Brown's paper on new Holland plants, under the name and disguise of Mr. Hibbert's gardener! Oh it is too bad!"; and James Edward Smith
James Edward Smith
Sir James Edward Smith was an English botanist and founder of the Linnean Society.Smith was born in Norwich in 1759, the son of a wealthy wool merchant. He displayed a precocious interest in the natural world...

 wrote that he had a copy of Knight's paper "but shall not keep it—I mean hereafter not to notice it or any other of the author's productions." Brown himself wrote of Salisbury "I scarcely know what to think of him except that he stands between a rogue and a fool."

With the passage of time, some dissenting views have emerged. In 1870, Charles Babbington wrote that "Knight's Proteaceae is very curious if the best part of it was really carried away from a Linnean meeting and appropriated by Salisbury to tease R. Brown"; and other authors have since denied Salisbury's involvement. In 1985 David Mabberley
David Mabberley
Professor Dr. David John Mabberley, , is a botanist, educator and writer. Among his varied interests is the taxonomy of tropical plants, especially trees of the families Labiatae, Meliaceae and Rutaceae. He is perhaps best known for his plant dictionary The plant-book. A portable dictionary of the...

offered the following in Salisbury's defence: "If Salisbury was guilty, why did he do it? ... [N]o doubt Salisbury felt that Brown was encroaching on his territory, for he had been writing on, and studying, Proteaceae for some time, and here was the Librarian of Smith's Linnean Society dashing off a paper to get his names into Hortus kewensis, criticizing Salisbury and abandoning some of his names on the way. It seems not inconceivable, therefore, that Salisbury... would have made amendments to Knight's final manuscript in the light of Brown's remarks... and hurried it through the press to defeat the machinations of his enemy."

Publication history

This paper has seen a number of republications. In total it has been published four times:
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