1 Vinris

Glaucium Corniculatum Classification Essay

Kew Bulletin

Description:Kew Bulletin is an international peer-reviewed journal of plant taxonomy. It publishes original articles, short communications and reviews on the taxonomy, nomnecalture, phylogeny, systematics and floristics of vascular plants and fungi. Papers on palynology, cytology, anatomy, phytogeography and phytochemistry are also included if they have taxonomic implications. Each part is illustrated with high quality line drawings and photographs. The journal also features a book review and notices section. From 2008 the journal will be published by Springer Verlag on behalf of the Royal Botanic Gardens, Kew, where the editorial office is located. It is available in print and online. Four parts are published each year (c. 640 pp).

Coverage: 1946-2014 (Vol. 1, No. 1 - Vol. 69, No. 4)

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ISSN: 00755974

EISSN: 1874933X

Subjects: Science & Mathematics, Botany & Plant Sciences, Biological Sciences

Collections: Biological Sciences Collection, Ecology & Botany II Collection, Life Sciences Collection

THE NAMES OF PLANTS The Names of Plants is a handy, two-part reference book for the botanist and amateur gardener. The book begins by documenting the historical problems associated with an ever-increasing number of common names of plants and the resolution of these problems through the introduction of International Codes for both botanical and horticultural nomenclature. It also outlines the rules to be followed when plant breeders name a new species or cultivar of plant. The second part of the book comprises an alphabetical glossary of generic and specific plant names, and components of these, from which the reader may interpret the existing names of plants and construct new names. For the third edition, the book has been updated to include explanations of the International Codes for both Botanical Nomenclature (2000) and Nomenclature for Cultivated Plants (1995). The glossary has similarly been expanded to incorporate many more commemorative names.


David Gledhill Formerly Senior Lecturer, Department of Botany, University of Bristol and Curator of Bristol University Botanic Garden

   Cambridge, New York, Melbourne, Madrid, Cape Town, Singapore, São Paulo Cambridge University Press The Edinburgh Building, Cambridge  , United Kingdom Published in the United States of America by Cambridge University Press, New York www.cambridge.org Information on this title: www.cambridge.org/9780521818636 © Cambridge University Press 2002 This book is in copyright. Subject to statutory exception and to the provision of relevant collective licensing agreements, no reproduction of any part may take place without the written permission of Cambridge University Press. First published in print format 2002 - isbn-13 978-0-511-06141-7 eBook (NetLibrary) - isbn-10 0-511-06141-2 eBook (NetLibrary) - isbn-13 978-0-521-81863-6 hardback -  hardback isbn-10 0-521-81863-X - isbn-13 978-0-521-52340-0 paperback - isbn-10 0-521-52340-0 paperback Cambridge University Press has no responsibility for the persistence or accuracy of s for external or third-party internet websites referred to in this book, and does not guarantee that any content on such websites is, or will remain, accurate or appropriate. © Cambridge University Press 1985, 1989, 2002


Preface to the first edition


Preface to the third edition The nature of the problem

ix 1

The size of the problem


Towards a solution to the problem The rules of botanical nomenclature Family names

15 25 29

Generic names


Species names


Epithets commemorating people


Geographical epithets


Categories below the rank of species




Synonymy and illegitimacy

44 46

The International Code of Nomenclature for Cultivated Plants Graft chimaeras

Botanical terminology The glossary

The glossary Addendum to glossary Bibliography

52 54 55 58 312 323


Preface to the first edition

Originally entitled The naming of plants and the meanings of plant names, this book is in two parts. The first part has been written as an account of the way in which the naming of plants has changed with time and why the changes were necessary. It has not been the writer’s intention to dwell upon the more fascinating aspects of common names but rather to progress from these to the situation which exists today; in which the botanical and horticultural names of plants must conform to internationally agreed standards. The aim has been to produce an interesting text which is equally as acceptable to the amateur gardener as to the botanist. The temptation to make this a definitive guide to the International Code of Botanical Nomenclature was resisted since others have done this already and with great clarity. A brief comment on synonymous and illegitimate botanical names and a reference to recent attempts to accommodate the various traits and interests in the naming of cultivated plants was added after the first edition. The book had its origins in a collection of Latin plant names, and their meanings in English, which continued to grow by the year but which could never be complete. Not all plant names have meaningful translations. Some of the botanical literature gives full citation of plant names (and translations of the names, as well as common names). There are, however, many horticultural and botanical publications in which plant names are used in a casual manner, or are misspelled, or are given meanings or common names that are neither translations nor common (in the world-wide sense). There is also a tendency that may be part of modern language, to reduce names of garden plants to an abbreviated form (e.g. Rhodo for Rhododendron). Literal names such as Vogel’s Napoleona, for Napoleona vogelii, [vii]

provide only limited information about the plant. The dedication of the genus to Napoleon Bonaparte is not informative. Only by further search of the literature will the reader find that Theodor Vogel was the botanist to the 1841 Niger expedition and that he collected some 150 specimens during a rainy July fortnight in Liberia. One of those specimens, number 45, was a Napoleona that was later named for him as the type of the new species by Hooker and Planchon. To have given such information would have made the text very much larger. The author has compiled a glossary which should serve to translate the more meaningful and descriptive names of plants from anywhere on earth but which will give little information about many of the people and places commemorated in plant names. Their entries do little more than identify the persons for whom the names were raised and their period in history. The author makes no claim that the glossary is all-encompassing or that the meanings he has listed are always the only meanings that have been put upon the various entries. Authors of Latin names have not always explained the meanings of the names they have erected and, consequently, such names may have been given different meanings by subsequent writers.


Preface to the third edition

Since making the assumption, in the second edition, that genetic manipulation of the properties of plants might require new consideration of the ways in which they are to be named, GM has proceeded apace. Not only can the innate genetic material be re-ordered – in ways that nature would have rejected through their exposure to natural selection by the environment – but alien genetic material, from other organisms, can be introduced to give bizarre results. Arabidopsis thaliana has only 10 chromosomes and has been the plant of choice for cytologists and nucleic acid workers because of this. The twenty-first century sees its genetic code mapped and its 25,000 genes being examined individually to ascertain the ‘meaning of plant life’. From quite practical beginnings such as giving tomato fruits an extended keeping time, to esoteric developments such as building a luminescence gene from a jellyfish into a mouse, there is now a proposal to insert a gene from an electric eel into plants so that the plants can provide sources of electricity. This new ‘green revolution’ has an historical ring of familiarity about it! The new century has not yet brought universal consistency in accepting the botanical and the horticultural codes. Yet science is already seeking to move towards an international biodiversity code for the naming of everything. If one was to be facetious, one might observe that man is still at 6’s and 7’s in seeking an explanation of everything – and may well, in the end, find that the answer is 42! The study of whole organisms and their systematic relationships is an economically unrewarding pure science but an essential area of continuing investigation. If man is intent on producing genetically deviant life forms, the descent of these must be known and their names must reflect that descent. [ix]

The nature of the problem A rose: by any name?

Man’s highly developed constructive curiosity and his capacity for communication are two of the attributes distinguishing him from all other animals. Man alone has sought to understand the whole living world and things beyond his own environment and to pass his knowledge on to others. Consequently, when he discovers or invents something new he also creates a new word, or words, in order to be able to communicate his discovery or invention to others. There are no rules to govern the manner in which such new words are formed other than those of their acceptance and acceptability. This is equally true of the common, or vulgar or vernacular names of plants. Such names present few problems until communication becomes multilingual and the number of plants named becomes excessive. For example, the diuretic dandelion is easily accommodated in European languages. As the lion’s tooth, it becomes Lowenzahn, dent de lion, dente di leone. As piss-abed it becomes Pissenlit, piscacane, and piscialetto. When further study reveals that there are more than a thousand different kinds of dandelion throughout Europe, the formulation of common names for these is both difficult and unacceptable. Common plant names present language at its richest and most imaginative (welcome home husband however drunk you be, for the houseleek or Sempervivum; shepherd’s weather-glass, for scarlet pimpernel or Anagallis; meet her i’th’entry kiss her i’th’buttery, or leap up and kiss me, for Viola tricolor; touch me not, for the balsam Impatiens noli-tangere; mind your own business, or mother of thousands, for Soleirolia soleirolii; blood drop emlets, for Mimulus luteus). Local variations in common names are numerous and this is perhaps a reflection of the importance of plants in general conversation, in [1]

the kitchen and in herbalism throughout the country in bygone days. An often quoted example of the multiplicity of vernacular names is that of Caltha palustris, for which, in addition to marsh marigold, kingcup and May blobs, there are 90 other local British names (one being dandelion), as well as over 140 German and 60 French vernacular names. Common plant names have many sources. Some came from antiquity by word of mouth as part of language itself, and the passage of time and changing circumstances have obscured their meanings. Fanciful ideas of a plant’s association with animals, ailments and festivities, and observations of plant structures, perfumes, colours, habitats and seasonality have all contributed to their naming. So too have their names in other languages. English plant names have come from Arabic, Persian, Greek, Latin, ancient British, AngloSaxon, Norman, Low German, Swedish and Danish. Such names were introduced together with the spices, grains, fruit plants and others which merchants and warring nations introduced to new areas. Foreign names often remained little altered but some were transliterated in such a way as to lose any meaning which they may have had originally. The element of fanciful association in vernacular plant names often drew upon comparisons with parts of the body and with bodily functions (priest’s pintle for Arum maculatum, open arse for Mespilus germanicus and arse smart for Polygonum hydropiper). Some of these persist but no longer strike us as ‘vulgar’ because they are ‘respectably’ modified or the associations themselves are no longer familiar to us (Arum maculatum is still known as cuckoo pint (cuckoo pintle) and as wake robin). Such was the sensitivity to indelicate names that Britten and Holland, in their Dictionary of English Plant Names (1886), wrote ‘We have also purposely excluded a few names which though graphic in their construction and meaning, interesting in their antiquity, and even yet in use in certain counties, are scarcely suited for publication in a work intended for general


readers’. They nevertheless included the examples above. The cleaning up of such names was a feature of the Victorian period, during which our common plant names were formalized and reduced in numbers. Some of the resulting names are prissy (bloody cranesbill, for Geranium sanguineum, becomes blood-red cranesbill), some are uninspired (naked ladies or meadow saffron, for Colchicum autumnale, becomes autumn crocus) and most are not very informative. This last point is not of any real importance because names do not need to have a meaning or be interpretable. Primarily, names are mere ciphers which are easier to use than lengthy descriptions and yet, when accepted, they can become quite as meaningful. Within limits, it is possible to use one name for a number of different things but, if the limits are exceeded, this may cause great confusion. There are many common plant names which refer to several plants but cause no problem so long as they are used only within their local areas or when they are used to convey only a general idea of the plant’s identity. For example, Wahlenberg ia saxicola in New Zealand, Phacelia whitlavia in southern California, USA, Clitoria ternatea in West Africa, Campanula rotundifolia in Scotland and Endymion non-scriptus (formerly Scilla non-scripta and now Hyacinthoides non-scripta) in England are all commonly called bluebells. In each area, local people will understand others who speak of bluebells but in all the areas except Scotland the song ‘The Bluebells of Scotland’, heard perhaps on the radio, will conjure up a wrong impression. At least ten different plants are given the common name of cuckoo-flower in England, signifying only that they flower in spring at a time when the cuckoo is first heard. The problem of plant names and of plant naming is that common names need not be formed according to any rule and can change as language, or the user of language, dictates. If our awareness extended only to some thousands of ‘kinds’ of plants we could manage by giving them numbers but, as our awareness extends, more ‘kinds’ are recognized and for most purposes we find a need


to organize our thoughts about them by giving them names and by forming them into named groups. Then we have to agree with others about the names and the groups, otherwise communication becomes hampered by ambiguity. A completely coded numerical system could be devised but would have little use to the non-specialist, without access to the details of encoding. Formalized names provide a partial solution to the two opposed problems presented by vernacular names: multiple naming of a single plant and multiple application of a single name. The predominantly two-word structure of such formal names has been adopted in recent historic times in all biological nomenclature, especially in the branch which, thanks to Isidorus Hispalensis (560–636), Archbishop of Seville, whose ‘Etymologies’ was a vast encyclopaedia of ancient learning and was studied for 900 years, we now call botany. Of necessity, botanical names have been formulated from former common names but this does not mean that in the translation of botanical names we may expect to find meaningful names in common language. Botanical names, however, do represent a stable system of nomenclature which is usable by people of all nationalities and has relevancy to a system of classification. Since man became wise, he has domesticated both plants and animals and, for at least the past 300 years, has bred and selected an ever growing number of ‘breeds’, ‘lines’ or ‘races’ of these. He has also given them names. In this, man has accelerated the processes which, we think, are the processes of natural evolution and has created a different level of artificially sustained, domesticated organisms. The names given by the breeders of the plants of the garden and the crops of agriculture and arboriculture present the same problems as those of vernacular and botanical names. Since the second edition of this book was published, genetic manipulation of the properties of plants has proceeded apace. Not only has the innate genetic material of plants been re-ordered, but alien genetic material, from other organisms, even from other kingdoms, has been introduced to give bizarre results. The products are unnatural and [4]

have not faced selection in nature. Indeed, some may present problems should they interbreed with natural populations in the future. There is still a divide between the international bodies concerned with botanical and cultivated plant names and the commercial interests that are protected by legislation for trademarking new genetic and transgenic products.


The size of the problem ‘Man by his nature desires to know’ (Aristotle)

Three centuries before Christ, Aristotle of Stagira, disciple of Plato, wrote extensively and systematically of all that was then known of the physical and living world. In this monumental task, he laid the foundations of inductive reasoning. When he died, he left his writings and his teaching garden to one of his pupils, Theophrastus (c. 370–285 BC), who also took over Aristotle’s peripatetic school. Theophrastus’ writings on mineralogy and plants totalled 227 treatises, of which nine books of Historia Plantarum contain a collection of contemporary knowledge about plants and eight of De Causis Plantarum are a collection of his own critical observations, a departure from earlier philosophical approaches, and rightly entitle him to be regarded as the father of botany. These works were subsequently translated into Syrian, to Arabic, to Latin and back to Greek. He recognized the distinctions between monocotyledons and dicotyledons, superior and inferior ovaries in flowers, the necessity for pollination and the sexuality of plants but, although he used names for plants of beauty, use or oddity, he did not try to name everything. To the ancients, as to the people of earlier civilizations of Persia and China, plants were distinguished on the basis of their culinary, medicinal and decorative uses – as well as their supposed supernatural properties. For this reason, plants were given a name as well as a description. Theophrastus wrote of some 500 ‘kinds’ of plant which, considering that material had been brought back from Alexander the Great’s campaigns throughout Persia, as far as India, would indicate a considerable lack of discrimination. In Britain, we now recognize more than that number of different ‘kinds’ of moss. [6]

Four centuries later, about AD 64, Dioscorides recorded 600 ‘kinds’ of plants and, half a century later still, the elder Pliny, in his huge compilation of the information contained in the writings of 473 authors, described about a thousand ‘kinds’. During the ‘Dark Ages’, despite the remarkable achievements of such people as Albertus Magnus (1193–1280), who collected plants during extensive journeys in Europe, and the publication of the German Herbarius in 1485 by another collector of European plants, Dr Johann von Cube, little progress was made in the study of plants. It was the renewal of critical observation by Renaissance botanists such as Dodoens (1517–1585), l’Obel (1538–1616), l’Ecluse (1526– 1609) and others which resulted in the recognition of some 4,000 ‘kinds’ of plants by the sixteenth century. At this point in history, the renewal of critical study and the beginning of plant collection throughout the known world produced a requirement for a rational system of grouping plants. Up to the sixteenth century, three factors had hindered such classification. The first of these was that the main interested parties were the nobility and apothecaries who conferred on plants great monetary value, either because of their rarity or because of the real or imaginary virtues attributed to them, and regarded them as items to be guarded jealously. Second was the lack of any standardized system of naming plants and third, and perhaps most important, any expression of the idea that living things could have evolved from earlier extinct ancestors and could therefore form groupings of related ‘kinds’ was a direct contradiction of the religious dogma of Divine Creation. Perhaps the greatest disservice to progress was that caused by the Doctrine of Signatures, which claimed that God had given to each ‘kind’ of plant some feature which could indicate the uses to which man could put the plant. Thus, plants with kidney-shaped leaves could be used for treating kidney complaints and were grouped together on this basis. Theophrastus Bombast von Hohenheim (1493– 1541) had invented properties for many plants under this doctrine. He also considered that man possessed intuitive knowledge of which [7]

plants could serve him, and how. He is better known under the Latin name which he assumed, Paracelsus, and the doctrinal book Dispensatory is usually attributed to him. The doctrine was also supported by Giambattista Della Porta (1543–1615), who made an interesting extension to it, that the distribution of different ‘kinds’ of plants had a direct bearing upon the distribution of different kinds of ailment which man suffered in different areas. On this basis, the preference of willows for wet habitats is ordained by God because men who live in wet areas are prone to suffer from rheumatism and, since the bark of Salix species gives relief from rheumatic pains (it contains salicylic acid, the analgesic principal of aspirin), the willows are there to serve the needs of man. In spite of disadvantageous attitudes, renewed critical interest in plants during the sixteenth century led to more discriminating views as to the nature of ‘kinds’, to searches for new plants from different areas and concern over the problems of naming plants. John Parkinson (1569–1629), a London apothecary, wrote a horticultural landmark with the punning title Paradisi in Sole -- Paradisus Terrestris of 1629. This was an encyclopaedia of gardening and of plants then in cultivation and contains a lament by Parkinson that, in their many catalogues, nurserymen ‘without consideration of kind or form, or other special note give(th) names so diversely one from the other, that . . . very few can tell what they mean’. This attitude towards common names is still with us but not in so violent a guise as that shown by an unknown author who, in Science Gossip of 1868, wrote that vulgar names of plants presented ‘a complete language of meaningless nonsense, almost impossible to retain and certainly worse than useless when remembered – a vast vocabulary of names, many of which signify that which is false, and most of which mean nothing at all’. Names continued to be formed as phrase-names constructed with a starting noun (which was later to become the generic name) followed by a description. So, we find that the creeping buttercup


was known by many names, of which Caspar Bauhin (1550–1624) and Christian Mentzel (1622–1701) listed the following: Caspar Bauhin, Pinax Theatri Botanici, 1623: Ranunculus pratensis repens hirsutus var. C.Bauhin repens fl. luteo simpl. J.Bauhin repens fol. ex albo variis repens magnus hirsutus fl. pleno repens flore pleno pratensis repens Parkinson pratensis reptante cauliculo l’Obel polyanthemos 1 Dodoens hortensis 1 Dodoens vinealis Tabernamontana pratensis etiamque hortensis Gerard Christianus Mentzelius, Index Nominum Plantarum Multilinguis (Universalis), 1682: Ranunculus pratensis et arvensis C.Bauhin rectus acris var. C.Bauhin rectus fl. simpl. luteo J.Bauhin rectus fol. pallidioribus hirsutis J.Bauhin albus fl. simpl. et denso J.Bauhin pratensis erectus dulcis C.Bauhin Ranoncole dolce Italian Grenoillette dor´ee o doux Gallic Sewite Woode Crawe foet English Suss Hanenfuss Jaskien sodky Polish Chrysanth. simplex Fuchs Ranunculus pratensis repens hirsutus var. c C.Bauhin repens fl. luteo simpl. J.Bauhin repens fol. ex albo variis Antonius Vallot repens magnus hirsut. fl. pleno J.B.Tabernamontana


repens fl. pleno J.Bauhin arvensis echinatus Paulus Ammannus prat. rad. verticilli modo rotunda C.Bauhin tuberosus major J.Bauhin Crus Galli Otto Brunfelsius Coronopus parvus Batrachion Apuleius Dodonaeus (Dodoens) Ranunculus prat. parvus fol. trifido C.Bauhin arvensis annuus fl. minimo luteo Morison fasciatus Henricus Volgnadius Ol. Borrich Caspar Bartholino These were, of course, common or vernacular names with wide currency and strong candidates for inclusion in lists which were intended to clarify the complicated state of plant naming. Local, vulgar names escaped such listing until much later times, when they were being less used and lexicographers began to collect them, saving most from vanishing for ever. Great advances were made during the seventeenth century. Robert Morison (1620–1683) published a convenient or artificial system of grouping ‘kinds’ into groups of increasing size, as a hierarchy. One of his groups we now call the family Umbelliferae or, to give it its modern name, Apiaceae, and this was the first natural group to be recognized. By natural group we imply that the members of the group share a sufficient number of common features to suggest that they have all evolved from a common ancestral stock. Joseph Pitton de Tournefort (1656–1708) had made a very methodical survey of plants and had assorted 10,000 ‘kinds’ into 698 groups (or genera). The ‘kinds’ must now be regarded as the basic units of classification called species. Although critical observation of structural and anatomical features led to classification advancing beyond the vague herbal and signature systems, no such advance was made in plant naming until a Swede, of little academic ability when young, we are told, established landmarks in both classification and nomenclature of plants. He was Carl Linnaeus (1707–1778), who classified [10]

7,300 species into 1,098 genera and gave to each species a binomial name (a name consisting of a generic name-word plus a descriptive epithet, both of Latin form). It was inevitable that, as man grouped the ever-increasing number of known plants (and he was then principally aware of those from Europe, the Mediterranean and a few from other areas) the constancy of associated morphological features in some groups should suggest that the whole was derived, by evolution, from a common ancestor. Morison’s family Umbelliferae was a case in point. Also, because the basic unit of any system of classification is the species, and some species were found to be far less constant than others, it was just as inevitable that the nature of the species itself would become a matter of controversy, not least in terms of religious dogma. A point often passed over with insufficient comment is that Linnaeus’ endeavours towards a natural system of classification were accompanied by his changing attitude towards Divine Creation. From the 365 aphorisms by which he expressed his views in Fundamenta Botanica (1736), and expanded in Critica Botanica (1737), his early view was that all species were produced by the hand of the Almighty Creator and that ‘variations in the outside shell’ were the work of ‘Nature in a sporty mood’. In such genera as Thalictrum and Clematis, he later concluded that some species were not original creations and, in Rosa, he was drawn to conclude that either some species had blended or that one species had given rise to several others. Later, he invoked hybridization as the process by which species could be created and attributed to the Almighty the creation of the primeval genera, each with a single species. From his observation of land accretion dur¨ ing trips to Oland and Gotland, in 1741, he accepted a continuous creation of the earth and that Nature was in continuous change (Oratio de Telluris habitabilis incremento, 1744). He later accepted that fossil bed remains could only be explained by a process of continuous creation. In Genera Plantarum, 6th edn. (1764), he attributed to God the creation of the natural orders (our families). Nature produced from these the genera and species, and permanent varieties [11]

were produced by hybridization between them. The abnormal varieties of the species so formed were the product of chance. Linnaeus was well aware of the results which plant hybridizers were obtaining in Holland and it is not surprising that his own knowledge of naturally occurring variants led him towards a covertly expressed belief in evolution. However, that expression, and his listing of varieties under their typical species in Species Plantarum, where he indicated each with a Greek letter, was still contrary to the dogma of Divine Creation and it would be another century before an authoritative declaration of evolutionary theory was to be made, by Charles Darwin (1809–1882). Darwin’s essay on ‘The Origin of Species by Means of Natural Selection’ (1859) was published somewhat reluctantly and in the face of fierce opposition. It was concerned with the major evolutionary changes by which species evolve and was based upon Darwin’s own observations on fossils and living creatures. The concept of natural selection, or the survival of any life-form being dependent upon its ability to compete successfully for a place in nature, became, and still is, accepted as the major force directing an inevitable process of organic change. Our conception of the mechanisms and the causative factors for the large evolutionary steps, such as the demise of the dinosaurs and of many plant groups now known only as fossils, and the emergence and diversification of the flowering plants during the last 100 million years is, at best, hazy. The great age of plant-hunting, from the second half of the eighteenth century through most of the nineteenth century, produced a flood of species not previously known. Strange and exotic plants were once prized above gold and caused theft, bribery and murder. Trading in ‘paper tulips’ by the van Bourse family gave rise to the continental stock exchange – the Bourse. With the invention of the Wardian Case by Dr Nathaniel Bagshaw Ward, in 1827, it became possible to transport plants from the farthest corners of the world by sea and without enormous losses. The case was a small glasshouse, which reduced water losses and made it unnecessary to [12]

use large quantities of fresh water on the plants during long sea voyages, as well as giving protection from salt spray. In the confusion which resulted from the naming of this flood of plants, and the use of many languages to describe them, it became apparent that there was a need for international agreement on both these matters. Today, we have rules formulated to govern the names of about 300,000 species of plants, which are now generally accepted, and have disposed of a great number of names that have been found invalid. Our present state of knowledge about the mechanisms of inheritance and change in plants and animals is almost entirely limited to an understanding of the causes of variation within a species. That understanding is based upon the observed behaviour of inherited characters as first recorded in Pisum by Gregor Johann Mendel, in 1866. With the technical development of the microscope, Malpighi (1671), Grew (1672) and others explored the cellular structure of plants and elucidated the mechanism of fertilization. However, the nature of inheritance and variability remained clouded by myth and monsters until Mendel’s work was rediscovered at the beginning of the twentieth century. By 1900, deVries, Correns, Tschermak and Bateson had confirmed that inheritance had a definite, particulate character which is regulated by ‘genes’. Sutton (1902) was the first person to clarify the manner in which the characters are transmitted from parents to offspring when he described the behaviour of ‘chromosomes’ during division of the cell nucleus. Chromosomes are thread-like bodies which can be stained in dividing cells so that the sequence of events of their own division can be followed. Along their length, it can be shown, the sites of genetic control, or genes, are situated in an ordered linear sequence. Differences between individuals can now be explained in terms of the different forms, or allelomorphs, in which single genes can exist as a consequence of their mutation. At the level of the gene, we must now consider the mutants and alleles as variants in molecular structure represented by the sequences of bases in the desoxyribonucleic acid. Classification [13]

can not yet accommodate the new, genetically modified forms that may only be distinguished in terms of some property resultant upon the insertion of a fragment of DNA. The concept of a taxonomic species, or grouping of individuals each of which has a close resemblance to the others in every aspect of its morphology, and to which a name can be applied, is not always the most accurate interpretation of the true circumstances in nature. It defines and delimits an entity but we are constantly discovering that the species is far from being an immutable entity. The botanist discovers that a species has components which have well-defined, individual ecotypic properties (an ability to live on a distinctive soil type, or an adaptation to flower and fruit in harmony with some agricultural practice) or have reproductive barriers caused by differences in chromosome number, etc. The plant breeder produces a steady stream of new varieties of cultivated species by hybridization and selection from the progeny. Genetically modified plants with very specific ‘economic’ properties are produced by techniques which evade nature’s safeguards of incompatibility and hybrid sterility and may or may not have to be repeatedly re-synthesized. If we consider some of the implications of, and attitudes towards, delimiting plant species and their components, and naming them, it will become easier to understand the need for internationally accepted rules intended to prevent the unnecessary and unacceptable proliferation of names.


Towards a solution to the problem

It is basic to the collector’s art to arrange items into groups. Postage stamps can be arranged by country of origin and then on face value, year of issue, design, colour variation, or defects. The arranging process always resolves into a hierarchic set of groups. In the plant kingdom we have a descending hierarchy of groups through Divisions, divided into Classes, divided into Orders, divided into Families, divided into Genera, divided into Species. Subsidiary groupings are possible at each level of this hierarchy and are employed to rationalize the uniformity of relationships within the particular group. Thus, a genus may be divided into a mini-hierarchy of subgenera, divided into sections, divided into series in order to assort the components into groupings of close relatives. All such components would, nevertheless, be members of the one genus. Early systems of classification were much less sophisticated and were based upon few aspects of plant structure such as those which suggested signatures, and mainly upon ancient herbal and medicinal concepts. Later systems would reflect advances in man’s comprehension of plant structure and function, and employ the morphology and anatomy of reproductive structures as defining features. Groupings such as Natural Orders and Genera had no precise limits or absolute parity, one with another; and genera are still very diverse in size, distribution and the extent to which they have been subdivided. Otto Brunfels (1489–1534) was probably the first person to introduce accurate, objective recording and illustration of plant structure in his Herbarium of 1530, and Valerius Cordus (1515– 1544) could have revolutionized botany but for his premature death. His four books of German plants contained detailed accounts of the structure of 446 plants, based upon his own systematic studies on [15]

them. Many of the plants were new to science. A fifth book on Italian plants was in compilation when he died. Conrad Gesner (1516–1565) published Cordus’ work on German plants in 1561 and the fifth book in 1563. A primitive suggestion of an evolutionary sequence was contained in Matthias de l’Obel’s Plantarum seu Stirpium Historia (1576) in which narrow-leaved plants, followed by broader-leaved, bulbous and rhizomatous plants, followed by herbaceous dicotyledons, followed by shrubs and trees, was regarded as a series of increasing ‘perfection’. Andrea Caesalpino (1519–1603) retained the distinction between woody and herbaceous plants but employed more detail of flower, fruit and seed structure in compiling his classes of plants (De Plantis, 1583). His influence extended to the classifications of Caspar Bauhin (1550–1624), who departed from the use of medicinal information and compiled detailed descriptions of the plants to which he gave many two-word names, or binomials. P.R. de Belleval (1558–1632) adopted a binomial system which named each plant with a Latin noun followed by a Greek adjectival epithet. Joachim Jung (1587–1657) feared being accused of heresy, which prevented him from publishing his work. The manuscripts which survived him contain many of the terms which we still use in describing leaf and flower structure and arrangement, and also contain plant names consisting of a noun qualified by an adjective. Robert Morison (1620–1683) used binomials, and John Ray (1627–1705), who introduced the distinction between monocotyledons and dicotyledons, but retained the distinction between flowering herbaceous plants and woody plants, also used binomial names. Joseph Pitton de Tournefort (1656–1708) placed great emphasis on the floral corolla and upon defining the genus, rather than the species. His 698 generic descriptions are detailed but his species descriptions are dependent upon binomials and illustrations. Herman Boerhaave (1668–1739) combined the systems of Ray and Tournefort, and others, to incorporate morphological, ecological, [16]

leaf, floral and fruiting characters, but none of these early advances received popular support. As Michel Adanson (1727–1806) was to realize, some sixty systems of classification had been proposed by the middle of the eighteenth century and none had been free from narrow conceptual restraints. His plea that attention should be focused on ‘natural’ classification through processes of inductive reasoning, because of the wide range of characteristics then being employed, did not enjoy wide publication and his work was not well regarded when it did become more widely known. His main claim to fame, or notoriety, stems from his use of names which have no meanings. Before considering the major contributions made by Carl Linnaeus, it should be noted that the names of many higher groups of plants, of families and of genera were well established at the beginning of the eighteenth century and several people had used simplified, binomial names for species. Indeed, August Quirinus Rivinus (1652–1723) had proposed that no plant should have a name of more than two words. Carl Linnaeus (1707–1778) was the son of a clergyman, Nils, who had adopted the latinized family name when he became a student of theology. Carl also went to theological college for a year but then left and became an assistant gardener in Prof. Olof Rudbeck’s botanic garden at Uppsala. His ability as a collector and arranger soon became evident and, after undertaking tours through Lapland, he began to publish works which are now the starting points for naming plants and animals. In literature he is referred to as Carl or Karl or Carolus Linnaeus, Carl Linn´e (an abbreviation) and, later in life, as Carl von Linn´e. His life became one of devotion to the classification and naming of all living things and of teaching others about them. His numerous students played a very important part in the discovery of new plants from many parts of the world. Linnaeus’ main contribution to botany was his method of naming plants, in which he combined Bauhin’s and Belleval’s use of binomials with Tournefort’s and Boerhaave’s concepts of the genus. His success, [17]

where others before him had failed, was due to the early publication of his most popular work, an artificial system of classifying plants. In this he employed the number, structure and disposition of the stamens of the flower to define 23 classes, each subdivided into orders on the basis of the number of parts constituting the pistil, with a 24th class containing those plants which had their reproductive organs hidden to the eye: the orders of which were the ferns, mosses, algae (in which he placed liverworts, lichens and sponges), fungi and palms. This ‘sexual system’ provided an easy way of grouping plants and of allocating newly discovered plants to a group. Originally designed to accommodate the plants of his home parish, it was elaborated to include first the Arctic flora and later the more diverse and exotic plants being discovered in the tropics. It continued in popular use into the nineteenth century despite its limitation of grouping together strange bedfellows: red valerian, tamarind, crocus, iris, galingale sedge and mat grass are all grouped under Triandria (three stamens) Monogynia (pistil with a single style). In 1735, Linnaeus published Systema Naturae, in which he grouped species into genera, genera into orders and orders into classes on the basis of structural similarities. This was an attempt to interpret evolutionary relationships or assemblages of individuals at different levels. It owed much to a collaborator and fellow student of Linnaeus, Peter Artendi (d. 1735) who, before an untimely death, was working on the classification of fishes, reptiles and amphibians, and the Umbelliferae. In Species Plantarum, published in 1753, Linnaeus gave each species a binomial name. The first word of each binomial was the name of the genus to which the species belonged and the second word was a descriptive, or specific epithet. Both words were in Latin or Latin form. Thus, the creeping buttercup he named as Ranunculus repens. It now required that the systematic classification and the binomial nomenclature, which Linnaeus had adopted, should become generally accepted and, largely because of the popularity of his sexual system, this was to be the case. Botany could now contend with [18]

the rapidly increasing number of species of plants being collected for scientific enquiry, rather than for medicine or exotic gardening, as in the seventeenth century. For the proper working of such standardized nomenclature, however, it was necessary that the language of plant names should also be standardized. Linnaeus’ views on the manner of forming plant names, and the use of Latin for these and for the descriptions of plants and their parts, have given rise directly to modern practice and a Latin vocabulary of great versatility, but which would have been largely incomprehensible in ancient Rome. He applied the same methodical principles to the naming of animals, minerals and diseases and, in doing so, established Latin, which was the lingua franca of his day, as the internationally used language of science and medicine. The rules by which we now name plants depend largely on Linnaeus’ writings but, for the names of plant families, we are much dependent on A.L. de Jussieu’s classification in his Genera Plantarum of 1789. For the name of a species, the correct name is that which was first published since 1753. This establishes Linnaeus’ Species Plantarum (associated with his Genera Plantarum, 5th edn. 1754 and 6th edn. 1764) as the starting point for the names of species (and their descriptions). Linnaeus’ sexual system of classification was very artificial and, although Linnaeus must have been delighted at its popularity, he regarded it as no more than a convenient pigeonholing system. He published some of his views on grouping plant genera into natural orders (our families) in Philosophia Botanica (1751). Most of his orders were not natural groupings but considerably mixed assemblages. By contrast, Bernard de Jussieu (1699–1777), followed by his nephew Antoine Laurent de Jussieu (1748–1836), searched for improved ways of arranging and grouping plants as natural groups. In A.L. de Jussieu’s de Genera Plantarum (1789) the characteristics are given for 100 plant families; and most of these we still recognize. Augustin Pyrame de Candolle (1778–1841) also sought a natural system, as did his son Alphonse, and he took the evolutionist view [19]

that there is an underlying state of symmetry in the floral structure which we can observe today and that, by considering relationships in terms of that symmetry, natural alliances may be recognized. This approach resulted in a great deal of monographic work from which de Candolle formed views on the concept of a core of similarity, or type, for any natural group and the requirement for control in the naming of plants. Today, technological and scientific advances have made it possible for us to use subcellular, chemical and the minutest of morphological features and to incorporate as many items of information as are available about a plant in computer-aided assessments of that plant’s relationships to others. Biological information has often been found to conflict with the concept of the taxonomic species and there are many plant groups in which the ‘species’ can best be regarded as a collection of highly variable populations. The gleaning of new evidence necessitates a continuing process of reappraisal of families, genera and species. Such reappraisal may result in subdivision or even splitting of a group into several new ones or, the converse process, in lumping together two or more former groups into one new one. Since the bulk of research is carried out on the individual species, most of the revisions are carried out at or below the rank of species. On occasion, therefore, a revision at the family level will require the transfer of whole genera from one family to another, but it is now more common for a revision at the level of the genus to require the transfer of some, if not all the species from one genus to another. Such revisions are not mischievous but are the necessary process by which newly acquired knowledge is incorporated into a generally accepted framework. It is because we continue to improve the extent of our knowledge of plants that revision of the systems for their classification continues and, consequently, that name changes are inevitable. The equivalence, certainly in evolutionary terms, of groups of higher rank than of family is a matter of philosophical debate and, even at the family level, we find divergence of views as to [20]

whether those with few components are equivalent to those with many components. Over the past twenty years a ‘Family Planning’ committee of taxonomists has met in London to determine an acceptable system of plant families in view of the variation presented by systematists since Bentham and Hooker. In the petaloid monocotyledons they were unanimous in agreeing to split the lilies (essentially the familiar families Liliaceae and Amaryllidaceae) to make the family concept more comparable with that adopted in other groups. The following liliaceous family names are now in common use: Melanthiaceae, Colchicaceae, Asphodelaceae, Hyacinthaceae, Hemerocallidaceae, Agavaceae, Aphyllandraceae, Lomandraceae, Anthericaceae, Xanthorrhoeaceae, Alliaceae, Liliaceae, Dracaenaceae, Asparagaceae, Ruscaceae, Convallariaceae, Trilliaceae, Alteriaceae, Herreriaceae, Philesiaceae, Smilacaceae, Haemadoraceae, Hypoxidaceae, Alstoemeriaceae, Doryanthaceae, Campynemaceae and Amaryllidaceae. Because the taxonomic species is the basic unit of any system of classification, we have to assume parity between species; that is to say, we assume that a widespread species is in every way comparable with a rare species which may be restricted in its distribution to a very small area. It is a feature of plants that their diversity – of habit, longevity, mode of reproduction and tolerance of environmental conditions – presents a wide range of biologically different circumstances. For the taxonomic problem of delimiting, defining and naming a species we have to identify a grouping of individuals whose characteristics are sufficiently stable to be defined, in order that a name can be applied to the group and a ‘type’, or exemplar, can be specified for that name. It is because of this concept of the ‘type’ that changes have to be made in names of species in the light of new discoveries and that entities below the rank of species have to be recognized. Thus, we speak of a botanical ‘sub-species’ when part of the species grouping can be distinguished as having a number of features which remain constant and as having a distinctive geographical or ecological distribution. When the degree of departure from the typical material is of a lesser order we may employ [21]

the inferior category of ‘variety’. The term ‘form’ is employed to describe a variant which is distinct in a minor way only, such as a single feature difference which might appear sporadically due to genetic mutation or sporting. The patterns and causes of variation differ from one species to another and this has long been recognized as a problem in fully reconciling the idea of a taxonomic species with that of a biological system of populations in perpetual evolutionary flux. Below the level of species, agreement about absolute ranking is far from complete and even the rigidity of the infraspecific hierarchy (subspecies, varietas, subvarietas, forma, subforma) is now open to question. It is always a cause of annoyance when a new name has to be given to a plant which is widely known under its superseded old name. Gardeners always complain about such name changes but there is no novelty in that. On the occasion of Linnaeus being proposed for Fellowship of the Royal Society, Peter Collinson wrote to him in praise of his Species Plantarum but, at the same time, complained that Linnaeus had introduced new names for so many well-known plants. The gardener has some cause to be aggrieved by changes in botanical names. Few gardeners show much alacrity in adopting new names and perusal of gardening books and catalogues shows that horticulture seldom uses botanical names with all the exactitude which they can provide. Horticulture, however, not only agreed to observe the international rules of botanical nomenclature but also formulated its own additional rules for the naming of plants grown under cultivation. It might appear as though the botanist realizes that he is bound by the rules, whereas the horticulturalist does not, but to understand this we must recognize the different facets of horticulture. The rules are of greatest interest and importance to specialist plant breeders and gardeners with a particular interest in a certain plant group. For the domestic gardener it is the growing of beautiful plants which is the motive force behind his activity. Between the two extremes lies every shade of interest and the main [22]

emphasis on names is an emphasis on garden names. Roses, cabbages, carnations and leeks are perfectly adequate names for the majority of gardeners but if greater precision is needed, a gardener wishes to know the name of the variety. Consequently, most gardeners are satisfied with a naming system which has no recourse to the botanical rules whatsoever. Not surprisingly, therefore, seed and plant catalogues also avoid botanical names. The specialist plant breeder, however, shows certain similarities to the apothecaries of an earlier age. Like them, he guards his art and his plants jealously because they represent the source of his future income and, also like them, he has the desire to understand every aspect of his plants. The apothecaries gave us the first centres of botanical enquiry and the plant breeders of today give us the new varieties which are needed to satisfy our gardening and food-production requirements. The commercial face of plant breeding, however, attaches a powerful monetary significance to the names given to new varieties. Gardeners occasionally have to resort to botanical names when they discover some cultural problem with a plant which shares the same common name with several different plants. The Guernsey lily, around which has always hung a cloud of mystery, has been offered to the public in the form of Amaryllis belladonna L. The true Guernsey lily has the name Nerine sarniensis Herb. (but was named Amaryllis sarniensis by Linnaeus). The epithet sarniensis means ‘of Sarnia’ or ‘of Guernsey’, Sarnia being the old name for Guernsey, and is an example of a misapplied geographical epithet, since the plant’s native area is South Africa. Some would regard the epithet as indicating the fact that Guernsey was the first place in which the plant was cultivated. This is historically incorrect, however, and does nothing to help the gardener who finds that the Guernsey lily that he has bought does not behave, in culture, as Nerine sarniensis is known to behave. This example is one involving a particularly contentious area as to the taxonomic problems of generic boundaries and typification but there are many others in which common and Latin garden names are used for whole assortments of garden plants, ranging [23]

from species (Nepeta mussinii and N. cataria are both catmint) to members of different genera (‘japonicas’ including Chaenomeles speciosa and Kerria japonica) to members of different families (Camellia japonica is likewise a ‘japonica’), and the diversity of ‘bluebells’ was mentioned earlier. New varieties, be they timber trees, crop plants or garden flowers, require names and those names need to be definitive. As with the earlier confusion of botanical names (different names for the same species or the same name for different species), so there can be the same confusion of horticultural names. As will be seen, rules for cultivated plants require that new names have to be established by publication. This gives to the breeder the commercial advantage of being able to supply to the public his new variety under what, initially, amounts to his mark of copyright. In some parts of the world legislation permits exemption from the rules and recommendations otherwise used for the names of cultivated plants.


The rules of botanical nomenclature

The rules which now govern the naming and the names of plants really had their beginnings in the views of A.P. de Candolle as he expressed them in his Th´eorie El´ementaire de la Botanique (1813). There, he advised that plants should have names in Latin (or Latin form but not compounded from different languages), formed according to the rules of Latin grammar and subject to the right of priority for the name given by the discoverer or the first describer. This advice was found inadequate and, in 1862, the International Botanical Congress in London adopted control over agreements on nomenclature. Alphonse de Candolle (1806–1893), who was A.P. de Candolle’s son, drew up four simple ‘Lois’, or laws, which were aimed at resolving what threatened to become a chaotic state of plant nomenclature. The Paris International Botanical Congress of 1867 adopted the Lois, which were: 1 One plant species shall have no more than one name. 2 No two plant species shall share the same name. 3 If a plant has two names, the name which is valid shall be that which was the earliest one to be published after 1753. 4 The author’s name shall be cited, after the name of the plant, in order to establish the sense in which the name is used and its priority over other names. It can be seen from the above Lois that, until the nineteenth century, botanists frequently gave names to plants with little regard either to the previous use of the same name or to names that had already been applied to the same plant. It is because of this aspect that one often encounters the words sensu and non inserted before the name of an author, although both terms are more commonly [25]

used in the sense of taxonomic revision, and indicate that the name is being used ‘in the sense of’ or ‘not in the sense of’ that author, respectively. The use of Latin, as the language in which descriptions and diagnoses were written, was not universal in the nineteenth century and many regional languages were used in different parts of the world. A description is an account of the plant’s habit, morphology and periodicity whereas a diagnosis is an author’s definitive statement of the plant’s diagnostic features, and circumscribes the limits outside which plants do not pertain to that named species. A diagnosis often states particular ways in which the species differs from another species of the same genus. Before the adoption of Latin as the accepted language of botanical nomenclature, searching for names already in existence for a particular plant, and confirming their applicability, involved searching through multilingual literature. The requirement to use Latin was written into the rules by the International Botanical Congress in Vienna, in 1905. However, the American Society of Plant Taxonomists produced its own Code in 1947, which became known as the Brittonia edition of the Rules or the Rochester Code, and disregarded this requirement. Not until 1959 was international agreement achieved and then the requirement to use Latin was made retroactive to January 1st, 1935, the year of the Amsterdam meeting of the Congress. The rules are considered at each International Botanical Congress, formerly held at five-, and more recently at six-, yearly intervals during peacetime. The International Code of Botanical Nomenclature (first published as such in 1952) was formulated at the Stockholm Congress of 1950. In 1930, the matter of determining the priority of specific epithets was the main point at issue. The practice of British botanists had been to regard that epithet which was first published after the plant had been allocated to its correct genus as the correct name. This has been called the Kew Rule, but it was defeated in favour of the rule that now gives priority to the epithet that was the first to be published from the starting date of [26]

May 1st, 1753. Epithets which predate the starting point, but which were adopted by Linnaeus, are attributed to Linnaeus (e.g. Bauhin’s Alsine media, Ammi majus, Anagyris foetida and Galium rubrum and Dodoens’ Angelica sylvestris are examples of binomials nevertheless credited to Linnaeus). The 1959 International Botanical Congress in Montreal introduced the requirement under the Code, that for valid publication of a name of a family or any taxon of lower rank, the author of that name should cite a ‘type’ for the name and that this requirement should be retrospective to January 1st, 1958. The idea of a type goes back to A.P. de Candolle and it implies a representative collection of characteristics to which a name applies. The type in Botany is a nomenclatural type: it is the type for the name and the name is permanently attached to it or associated with it. For the name of a family, the representative characteristics which that name implies are those embodied in one of its genera, which is called the type genus. In a similar way, the type for the name of a genus is the type species of that genus. For the name of a species or taxon of lower rank, the type is a specimen lodged in an herbarium or, in certain cases, published illustrations. The type need not, nor could it, be representative of the full range of entities to which the name is applied. Just as a genus, although having the features of its parent family, cannot be fully representative of all the genera belonging to that family, no single specimen can be representative of the full range of variety found within a species. For the name to become the correct name of a plant or plant group, it must satisfy two sets of conditions. First, it must be constructed in accordance with the rules of name formation, which ensures its legitimacy. Second, it must be published in such a way as to make it valid. Publication has to be in printed matter which is distributed to the general public or, at least, to botanical institutions with libraries accessible to botanists generally. Since January 1st, 1953, this has excluded publication in newspapers and tradesmen’s [27]

catalogues. Valid publication also requires the name to be accompanied by a description or diagnosis, an indication of its rank and the nomenclatural type, as required by the rules. This publication requirement, and subsequent citation of the new name followed by the name of its author, ensures that a date can be placed upon the name’s publication and that it can, therefore, be properly considered in matters of priority. The present scope of the Code is expressed in the Principles, which have evolved from the de Candollean Lois: 1 Botanical nomenclature is independent of zoological nomenclature. The Code applies equally to names of taxonomic groups treated as plants whether or not these groups were originally so treated. 2 The application of names of taxonomic groups is determined by means of nomenclatural types. 3 The nomenclature of a taxonomic group is based upon priority of publication. 4 Each taxonomic group with a particular circumscription, position and rank can bear only one correct name, the earliest which is in accordance with the rules, except in specified cases. 5 Scientific names of taxonomic groups are treated as Latin, regardless of their derivation. 6 The rules of nomenclature are retroactive unless expressly limited. The detailed rules are contained in the Articles and Recommendations of the Code and mastery of these can only be gained by practical experience (Greuter, 2000). A most lucid summary and comparison with other Codes of biological nomenclature is that of Jeffrey (1978), written for the Systematics Association. There are still new species of plants to be discovered and an enormous amount of information yet to be sought for long-familiar species, in particular, evidence of a chemical nature, and especially that concerned with proteins, which may provide reliable indications of phylogenetic relationships. For modern systematists, the [28]

greatest and most persistent problem is our ignorance about the apparently explosive appearance of a diverse array of flowering plants, some 100 million years ago, from one or more unknown ancestors. Modern systems of classification are still frameworks within which the authors arrange assemblages in sequences or clusters to represent their own idiosyncratic interpretation of the known facts. In addition to having no firm record of the early evolutionary pathways of the flowering plants, the systematist also has the major problems of identifying clear-cut boundaries between groups and of assessing the absolute ranking of groups. It is because of these continuing problems that, although the Code extends to taxa of all ranks, most of the rules are concerned with the names and naming of groups from the rank of family downwards. Before moving on to the question of plant names at the generic and lower ranks, this is a suitable point at which to comment on new names for families which are now starting to appear in books and catalogues, and some explanation in passing may help to dispel any confusion. The splitting of the Liliaceae and Amaryllidaceae into 27 new families was mentioned on page 21 but the move towards standardization has required other family name changes.

Family names Each family can have only one correct name and that, of course, is the earliest legitimate one, except in cases of limitation of priority by conservation. In other words, there is provision in the Code for disregarding the requirement of priority when a special case is proved for a name to be conserved. Conservation of names is intended to avoid disadvantageous name changes, even though the name in question does not meet all the requirements of the Code. Names which have long-standing use and wide acceptability and are used in standard works of literature can be proposed for conservation and, when accepted, need not be discarded in favour of new and more correct names. [29]

The names of families are plural adjectives used as nouns and are formed by adding the suffix -aceae to the stem, which is the name of an included genus. Thus, the buttercup genus Ranunculus gives us the name Ranunculaceae for the buttercup family and the water-lily genus Nymphaea gives us the name Nymphaeaceae for the water-lilies. A few family names are conserved, for the reasons given above, which do have generic names as their stem, although one, the Ebenaceae, has the name Ebenus Kuntze (1891) non Linnaeus (1753) as its stem. Kuntze’s genus is now called Maba but its parent family retains the name Ebenaceae even though Ebenus L. is the name used for a genus of the pea family. There are eight families for which specific exceptions are provided and which can be referred to either by their long-standing, conserved names or, as is increasingly the case in recent floras and other published works on plants, by their names which are in agreement with the Code. These families and their equivalent names are: Compositae Cruciferae Gramineae Guttiferae Labiatae Leguminosae Palmae Umbelliferae

or or or or or or or or

Asteraceae (on the genus Aster) Brassicaceae (on the genus Brassica) Poaceae (on the genus Poa) Clusiaceae (on the genus Clusia) Lamiaceae (on the genus Lamium) Fabaceae (on the genus Faba) Arecaceae (on the genus Areca) Apiaceae (on the genus Apium)

Some botanists regard the Leguminosae as including three subfamilies but others accept those three components as each having family status. In the latter case, the three families are the Caesalpiniaceae, the Mimosaceae and the Papilionaceae. The last of these family names refers to the resemblance which may be seen in the pea- or bean-flower structure, with its large and colourful sail petal, to a resting butterfly (Papilionoidea) and is not based upon the name of a plant genus. If a botanist wishes to retain the three-family concept, the name Papilionaceae is conserved against Leguminosae and the [30]

modern equivalent is Fabaceae. Consequently, the Fabaceae are either the entire aggregation of leguminous plant genera or that part of the aggregate which does not belong in either the Caesalpiniaceae or the Mimosaceae. Some eastern European publications use Daucaceae for the Apiaceae, split the Asteraceae into Carduaceae and Chicoriaceae and adopt various views as to the generic basis of family names (e.g. Oenotheraceae for Onagraceae by insisting that Linnaeus’ genus Oenothera has prior claim over Miller’s genus Onagra).

Generic names The name of a genus is a noun, or word treated as such, and begins with a capital letter. It is singular, may be taken from any source whatever, and may even be composed in an arbitrary manner. The etymology of generic names is, therefore, not always complete and, even though the derivation of some may be discovered, they lack meaning. By way of examples: Portulaca, from the Latin porto (I carry) and lac (milk) translates as ‘Milk-carrier’. Pittosporum, from the Greek, pittow (I tar) and sporov (a seed) translates as ‘Tar-seed’. Hebe was the goddess of youth and, amongst other things, the daughter of Jupiter. It cannot be translated further. Petunia is taken from the Brazilian name for tobacco. Tecoma is taken from a Mexican name. Linnaea is one of the names which commemorate Linnaeus. Sibara is an anagram of Arabis. Aa is the name given by Reichenbach to an orchid genus which he segregated from Altensteinia. It has no meaning and, as others have observed, must always appear first in an alphabetic listing. The generic names of some Old World plants were taken from Greek mythology by the ancients, or are identical to the names of characters in Greek mythology. The reason for this is not always [31]

clear (e.g. Althaea, Cecropia, Circaea, Melia, Phoenix, Tagetes, Thalia, Endymion, Hebe, Paeonia and Paris). However, some do have reasonable floristic associations, e.g. Atropa (the third Fate, who held the scissors to cut the thread of life), Chloris (the Goddess of flowers), Iris (messenger to Gods of the rainbow), Melissa (apiarist who used the plant to feed the bees). The metamorphoses, that are so common in the mythology, provided direct associations for several names, e.g. Acanthus (became an Acanthus), Adonis (became an Anemone), Ajacis (became a Narcissus), Daphne (became a laurel), Hyacinthus (became, probably, a Delphinium) and Narcissus (became a daffodil). If all specific names were constructed in the arbitrary manner used by M. Adanson (1727–1806), there would have been no enquiries of the author and this book would not have been written. In fact, the etymology of plant names is a rich store of historical interest and conceals many facets of humanity ranging from the sarcasm of some authors to the humour of others. This is made possible by the wide scope available to authors for formulating names and because, whatever language is the source, names are treated as being in Latin. Imaginative association has produced some names which are very descriptive provided that the reader can spot the association. In the algae, the Chrysophyte which twirls like a ballerina has been named Pavlova gyrans and, in the fungi, a saprophyte on leaves of Eucalyptus which has a wide-mouthed spore-producing structure has been named Satchmopsis brasiliensis (Satchmo, satchelmouth). The large vocabulary of botanical Latin comes mostly from the Greek and Latin of ancient times but, since the ancients had few words which related specifically to plants and their parts, a Latin dictionary is of somewhat limited use in trying to decipher plant diagnoses. By way of examples, Table 1 gives the parts of the flower (Latin flos, Greek anqov) (illustrated in Fig. 1) and the classical words from which they are derived, together with their original sense. The grammar of botanical Latin is very formal and much more simple than that of the classical language itself. A full and most authoritative work on the subject is contained in Stearn’s book, [32]

Table 1 Flower part



Former meaning


kalux ´

k´ul´ix sk´eph — p´etalon — — — — audr-, ’ o’ik´ov st´igma stulov — karp´ov gun´h-, o’ik´ov —

— — corolla — petalum stamen filamentum anthera — — — stilus — — pistillum

various kinds of covering cup or goblet covering garland or coronet leaf metal plate thread, warp, string thread potion of herbs man-, house tattoo or spot pillar or post pointed writing tool fruit woman-, house pestle

sepal corolla petal stamen filament anther androecium stigma style carpel gynoecium pistil

Botanical Latin (1983). Nevertheless, it is necessary to know that in Latin, nouns (such as family and generic names) have gender, number and case and that the words which give some attribute to a noun (as in adjectival specific epithets) must agree with the noun in each of these. Having gender means that all things (the names of which are called nouns) are either masculine or feminine or neuter. In English, we treat almost everything as neuter, referring to nouns as ‘it’, except animals and most ships and aeroplanes (which are commonly held to be feminine). Gender is explained further below. Number means that things may be single (singular) or multiple (plural). In English we either have different words for the singular and plural (man and men, mouse and mice) or we convert the singular into the plural most commonly by adding an ‘s’ (ship and [33]

ships, rat and rats) or more rarely by adding ‘es’ (box and boxes, fox and foxes) or, rarer still, by adding ‘en’ (ox and oxen). In Latin, the difference is expressed by changes in the endings of the words. Case is less easy to understand but means the significance of the noun to the meaning of the sentence in which it is contained. It is also expressed in the endings of the words. In the sentence, ‘The flower has charm’, the flower is singular, is the subject of the sentence and has what is called the nominative case. In the sentence ‘I threw away the flower’, I am now the subject and the flower has become the direct object in the accusative case. In the sentence, ‘I did not like the colour of the flower’, I am again the subject, the colour is now the object and the flower has become a possessive noun and has the genitive case. In the sentence, ‘The flower fell to the ground’ the flower is once again the subject (nominative) and the ground has the dative case. If we add ‘with a whisper’, then whisper takes the ablative case. In other words, case confers on nouns an expression of their meaning in any sentence. This is shown by the ending of the Latin word, which changes with case and number and, in so doing, changes the naked word into part of a sentence (Table 2). Nouns fall into five groups, or declensions, as determined by their endings (Table 3). Generic names are treated as singular subjects, taking the nominative case. Solanum means ‘Comforter’ and derives from the use Table 2 Case


nominative accusative genitive dative ablative

flos the flower (subject) florem the flower (object) floris of the flower flori to or for the flower flore by, with or from the flower


Plural flores the flowers flores the flowers florum of the flowers floribus to or for the flowers floribus by, with or from the flowers


-a -am -ae -ae -a

nom acc gen dat abl

nom acc gen dat abl





-i -os -orum -is -is

-us(-er) -um -i -o -o



Denotes various irregular endings.

-ae -as -arum -is -is



Table 3

-a -a -orum -is -is

-um -um -i -o -o


-es -es -um -ibus -ibus

* -em -is -i -e



-a -a -um -ibus -ibus

* * -is -i -e


-es -es(is) -ium -ibus -ibus

-is(es) -em(im) -is -i -i(e)


-ia -ia -ium -ibus -ibus

-e(l)(r) -e(l)(r) -is -i -i(e)


-us -us -uum -ibus -ibus

-us -um -us -ui(u) -u



-ua -ua -uum -ibus -ibus

-u -u -us -ui(u) -u


-es -es -erum -ebus -ebus

-es -em -ei -ei -e



of nightshades as herbal sedatives. The gender of generic names is that of the original Greek or Latin noun or, if that was variable, is chosen by the author of the name. There are exceptions to this in which masculine names are treated as feminine, and fewer in which compound names, which ought to be feminine, are treated as masculine. As a general guide, names ending in -us are masculine unless they are trees (such as Fagus, Pinus, Quercus, Sorbus which are treated as feminine), names ending in -a are feminine and names ending in -um are neuter; names ending in -on are masculine unless they can also take -um, when they are neuter, or the ending is -dendron when they are also neuter (Rhododendron or Rhododendrum); names ending in -ma (as in terminations such as -osma) are neuter; names ending in -is are mostly feminine or masculine treated as feminine (Orchis) and those ending in -e are neuter; other feminine endings are -ago, -odes, -oides, -ix and -es. A recommendation for forming generic names to commemorate men or women is that these should be treated as feminine and formed as follows: for names ending in a vowel, for names ending in -a, for names ending in -ea, for names ending in a consonant, for names ending in -er, for latinized names ending in -us,

terminate with -a terminate with -ea do not change add -ia add -a change the ending to -ia

Generic names which are formed arbitrarily or are derived from vernacular names have their ending selected by the name’s author.

Species names The name of a species is a binary combination of the generic name followed by a specific epithet. If the epithet is of two words they must be joined by a hyphen or united into one word. The epithet can be taken from any source whatever and may be constructed [36]

Table 4 Masculine





-us -is -os -er -er -ax -ex -ox -ans -ens -or -oides

-a -is -os -era -ra -ax -ex -ox -ans -ens -or -oides

-um -e -on -erum -rum -ax -ex -ox -ans -ens -or -oides

hirsutus brevis acaulos Škaulov asper scaber fallax duplex ferox reptans repens tricolor bryoides brÅon, e«dov

(hairy) (short) (stemless) (rough) (rough) (false) (double) (very prickly) (creeping) (creeping) (three-coloured) (moss-like)

Table 5 Masculine





-us -ior -issimus -is -ior -limus -er -erior -errimus

-a -ior -issima -is -ior -lima -era -erior -errima

-um -ius -issimum -e -ius -limum -erum -erius -errimum


(long) (longer) (longest) (slender) (slenderer) (slenderest) (thin) (thinner) (thinnest)



in an arbitrary manner. It would be reasonable to expect that the epithet should have a descriptive purpose, and there are many which do, but large numbers either refer to the native area in which the plant grows or commemorate a person (often the discoverer, the [37]

introducer into cultivation or a noble personage). The epithet may be adjectival (or descriptive), qualified in various ways with prefixes and suffixes, or a noun. It will become clear that because descriptive, adjectival epithets must agree with the generic name, the endings must change in gender, case and number; Dipsacus fullonum L. has the generic name used by Dioscorides meaning ‘Dropsy’, alluding to the accumulation of water in the leaf-bases, and an epithet which is the masculine genitive plural of fullo, a fuller, and which identifies the typical form of this teasel as the one which was used to clean and comb up a ‘nap’ on cloth. The majority of adjectival epithet endings are as in the first two examples listed in Table 4. Comparative epithets are informative because they provide us with an indication of how the species contrasts with the general features of other members of the genus (Table 5).

Epithets commemorating people Specific epithets which are nouns are grammatically independent of the generic name. Campanula trachelium is literally ‘Little bell’ (feminine) ‘neck’ (neuter). When they are derived from the names of people, they can either be retained as nouns in the genitive case (clusii is the genitive singular of Clusius, the latinized version of l’Ecluse, and gives an epithet with the meaning ‘of l’Ecluse’) or be treated as adjectives and then agreeing in gender with the generic noun (Sorbus leyana Wilmott is a tree taking, like many others, the feminine gender despite the masculine ending, and so the epithet which commemorates Augustin Ley also takes the feminine ending). The epithets are formed as follows: to names ending with a vowel (except -a) or -er is added i when masculine singular, ae when feminine singular, orum when masculine plural, arum when feminine plural [38]

to names ending with -a is added e when singular, rum when plural to names ending with a consonant (except -er) is added ii when masculine singular, iae when feminine singular, iorum when masculine plural, iarum when feminine plural or, when used adjectivally: to names ending with a vowel (except -a) is added anus when masculine, ana when feminine, anum when neuter to names ending with -a is added nus when masculine, na when feminine, num when neuter to names ending with a consonant is added ianus when masculine, iana when feminine, ianum when neuter.

Geographical epithets When an epithet is derived from the name of a place, usually to indicate the plant’s native area but also, sometimes, to indicate the area or place from which the plant was first known or in which it was produced horticulturally, it is preferably adjectival and takes one of the following endings: -ensis (m) -(a)nus (m) -inus (m) -icus (m)

-ensis (f ) -(a)na (f ) -ina (f ) -ica (f )

-ense (n) -(a)num (n) -inum (n) -icum (n) [39]

Geographical epithets are sometimes inaccurate because the author of the name was in error as to the true origin of the plant, or obscure because the ancient classical names are no longer familiar to us. As with epithets which are derived from proper names to commemorate people, or from generic names or vernacular names which are treated as being Latin, it is now customary to start them with a small initial letter but it remains permissible to give them a capital initial.

Categories below the rank of species The subdivision of a species group is based upon a concept of infraspecific variation which assumes that, in nature, evolutionary changes are progressive fragmentations of the parent species. Put in another way, a species, or any taxon of lower rank, is a closed grouping whose limits embrace all their lower-ranked variants (subordinate taxa). It will be seen later that a different concept underlies the naming of cultivated plants which does not make such an assumption but recognizes the possibility that cultivars may straddle species, or other, boundaries or overlap each other, or be totally contained, one by another. The rules by which botanical infraspecific taxa are named specify that the name shall consist of the name of the parent species followed by a term which denotes the rank of the subdivision, and an epithet which is formed in the same ways as specific epithets, including grammatical agreement when adjectival. Such names are subject to the rules of priority and typification. The ranks concerned are subspecies (abbreviated to subsp. or ssp.), varietas (variety in English, abbreviated to var.), subvarietas (subvariety or subvar.), forma (form or f.). These form a hierarchy and further subdivisions are permitted but the Code does not define the characteristics of any rank within the hierarchy. Consequently, infraspecific classification is subjective.


When a subdivision of a species is named, which does not include the nomenclatural type of the species, it automatically establishes the name of the equivalent subdivision which does contain that type. Such a name is an ‘autonym’ and has the same epithet as the species itself but is not attributed to an author. This is the only event which permits the repetition of the specific epithet and the only permissible way of indicating that the taxon includes the type for the species name. The same constraints apply to subdivisions of lower ranks. For example, Veronica hybrida L. was deemed by E.F. Warburg to be a component of Veronica spicata L. and he named it V. spicata L. subsp. hybrida (L.) E.F. Warburg. This implies the existence of a typical subspecies, the autonym for which is V. spicata L. subsp. spicata. It will be seen from the citation of Warburg’s new combination that the disappearance of a former Linnaean species can be explained. Retention of the epithet ‘hybrida’, and the indication of Linnaeus being its author (in parentheses) shows the benefit of this system in constructing names with historic meanings.

Hybrids Hybrids are particularly important as cultivated plants but are also a feature of many plant groups in the wild, especially woody perennials such as willows. The rules for the names and naming of hybrids are contained in the Botanical Code but are equally applicable to cultivated plant hybrids. For the name of a hybrid between parents from two different genera, a name can be constructed from the two generic names, in part or in entirety (but not both in their entirety) as a condensed formula; × Mahoberberis is the name for hybrids between the genera Mahonia and Berberis (in this case the cross is only bigeneric when Mahonia, a name conserved against Berberis, is treated as a distinct genus) and × Fatshedera is the name for hybrids between the genera Fatsia and Hedera. The orchid hybrid between Gastrochilus bellinus


(Rchb.f.) O.Ktze. and Doritis pulcherrima Lindl. carries the hybrid genus name ×Gastritis (it has a cultivar called ‘Rumbling Tum’!). Alternatively a formula can be used in which the names of the genera are linked by the sign for hybridity ‘×’: Mahonia × Berberis and Fatsia × Hedera. Hybrids between parents from three genera are also named either by a formula or by a condensed formula and, in all cases, the condensed formula is treated as a generic name if it is published with a statement of parentage. When published, it becomes the correct generic name for any hybrids between species of the named parental genera. A third alternative is to construct a commemorative name in honour of a notable person and to end it with the termination -ara: × Sanderara is the name applied to the orchid hybrids between the genera Brassia, Cochlioda and Odontoglossum and commemorates H.F.C. Sander, the British orchidologist. A name formulated to define a hybrid between two particular species from different genera can take the form of a species name, and then applies to all hybrids produced subsequently from those parent species: × Fatshedera lizei Guillaumin is the name first given to the hybrid between Fatsia japonica (Thunb.) Decne. & Planch. and Hedera helix L. cv. Hibernica, but which must include all hybrids between F. japonica and H. helix; and × Cupressocyparis leylandii (Jackson & Dallimore) Dallimore is the name for hybrids between Chamaecyparis nootkatensis (D.Don) Spach and Cupressus macrocarpa Hartweg ex Godron. Other examples include × Achicodonia, × Achimenantha, × Amaryg ia, × Celsioverbascum, × Citrofortunella, × Chionoscilla, × Cooperanthes, × Halimocistus, × Ledodendron, × Leucoraoulia, × Lycene, × Osmarea, × Stravinia, × Smithicodonia, × Solidaster and × Venidioarctotis. Because the parents themselves are variable, the progeny of repeated crosses may be distinctive and warrant naming. They may be named under the Botanical Code (prior to 1982 they would have been referred to as nothomorphs or bastard forms) and also under the International Code of Nomenclature for Cultivated Plants as ‘cultivars’: thus, × Cupressocyparis leylandii cv. Naylor’s Blue. The hybrid nature of × Sanderara is expressed by [42]

classifying it as a ‘nothogenus’ (bastard genus or, in the special circumstances of orchid nomenclature, grex class) and of ×Cupressocyparis leylandii by classifying it as a ‘nothospecies’ (within a nothogenus). For infraspecific ranks the multiplication sign is not used but the term denoting their rank receives the prefix notho-, or ‘n-’ (Mentha × piperita L. nothosubspecies pyramidalis (Ten.) Harley which, as stated earlier, also implies the autonymous Mentha × piperita nothosubspecies piperita. Hybrids between species in the same genus are also named by a formula or by a new distinctive epithet: Dig italis lutea L. × D. purpurea L. and Nepeta × faassenii Bergmans ex Stearn are both correct designations for hybrids. In the example of Dig italis, the order in which the parents are presented happens to be the correct order, with the seed parent first. It is permissible to indicate the roles of the parents by including the symbols for female ‘乆’ and male ‘么’, when this information is known, or otherwise to present the parents in alphabetical order. The orchid family presents particularly complex problems of nomenclature, requiring its own ‘Code’ in the form of the Handbook on Orchid Nomenclature and Reg istration (Greatwood, Hunt, Cribb & Stewart, 1993). There are some 20,000 species of orchids and to this has been added a huge range of hybrids, some with eight genera contributing to their parentage, and over 70,000 hybrid swarms, or greges (singular grex–a crowd or troupe), with a highly complex ancestral history. In cases where a hybrid is sterile because the two sets of chromosomes which it has inherited, one from each parent, are sufficiently dissimilar to cause breakdown of the mechanism which ends in the production of gametes, doubling its chromosome complement may produce a new state of sexual fertility and what is, in effect, a new biological species. Many naturally occurring species are thought to have evolved by such changes and man has created others artificially via the same route, some intentionally and some unintentionally from the wild. The bread-wheats, Triticum aestivum L. [43]

are an example of the latter. They are not known in the wild and provide an example of a complex hybrid ancestry but whose name does not need to be designated as hybrid. Even artificially created tetraploids (having, as above, four instead of the normal two sets of chromosomes) need not be designated as hybrid, by inclusion of ‘×’ in the name: Dig italis mertonensis Buxton & Darlington is the tetraploid from an infertile hybrid between D. grandiflora L. and D. purpurea L.

Synonymy and illegitimacy Inevitably, most plants have been known by two or more names in the past. Since a plant can have only one correct name, which is determined by priority, its other validly published names are synonyms. A synonym may be one which is strictly referable to the same type (a nomenclatural synonym) or one which is referable to another type which is, however, considered to be part of the same taxon (this is a taxonomic synonym). The synonymy for any plant or group of plants is important because it provides a reference list to the history of the classification and descriptive literature on that plant or group of plants. In the search for the correct name, by priority, there may be names which have to be excluded from consideration because they are regarded as being illegitimate, or not in accordance with the rules. Names which have the same spelling but are based on different types from that which has priority are illegitimate ‘junior homonyms’. Clearly, this prevents the same name being used for different plants. Curiously, this exclusion also applies to the names of those animals which were once regarded as plants, but not to any other animal names. Published names of taxa, which are found to include the type of an existing name, are illegitimate because they are ‘superfluous’. This prevents unnecessary and unacceptable proliferation of names of no real value. [44]

Names of species in which the epithet exactly repeats the generic name have to be rejected as illegitimate ‘tautonyms’. It is interesting to note that there are many plant names which have achieved some pleonastic repetition by using generic names with Greek derivation and epithets with Latin derivation: Arctostaphylos uva-ursi (bearberry, berry of the bear); Myristica fragrans (smelling of myrrh, fragrant), Orobanche rapum-genistae (legume strangler, rape of broom); or the reverse of this, Liquidambar styraciflua (liquid amber, flowing with storax); Silaum silaus; but modern practice is to avoid such constructions. In zoological nomenclature tautonyms are commonplace. The Code provides a way of reducing unwelcome disturbance to customary usage which would be caused by rigid application of the rule of priority to replace with correct names certain names of families and genera which, although incorrect or problematic are, for various reasons (usually their long usage and wide currency in important literature) agreed to be conserved at a Botanical Congress. These conserved names can be found listed in an Appendix to the Code, together with names which are to be rejected because they are taxonomic synonyms used in a sense which does not include the type of the name, or are earlier nomenclatural synonyms based on the same type, or are homonyms or orthographic variants. The Code also recommends the ways in which names should be spelt or transliterated into Latin form in order to avoid what it refers to as ‘orthographic variants’. The variety found amongst botanical names includes differences in spelling which are, however, correct because their authors chose the spellings when they published them and differences which are not correct because they contain any of a range of defects which have become specified in the Code. This is a problem area in horticultural literature, where such variants are commonplace. It is clearly desirable that a plant name should have a single, constant and correct spelling but this has not been achieved in all fields and reaches its worst condition in the labelling of plants for sale in some nurseries. [45]

The International Code of Nomenclature for Cultivated Plants

There can be no doubt that the diverse approaches to naming garden plants, by common names, by botanical names, by mixtures of botanical and common names, by group names and by fancy names, is no less complex than the former unregulated use of common or vernacular names. The psychology of advertising takes descriptive naming into yet new dimensions. It catches the eye with bargain offers of colourful, vigorous and hardy, large-headed, incurved Chrysanthemum cvs. by referring to them as HARDY FOOTBALL MUMS. However, we are not here concerned with such colloquial names or the ethics of mail-order selling techniques but with the regulation of meaningful names under the Code. In 1952, the Committee for the Nomenclature of Cultivated Plants of the International Botanical Congress and the International Horticultural Congress in London adopted the International Code of Nomenclature for Cultivated Plants. Sometimes known as the Cultivated Code, it was first published in 1953 and has been revised several times at irregular intervals since then (Trehane, 1995). This Code formally introduced the term ‘cultivar’ to encompass all varieties or derivatives of wild plants which are raised under cultivation and its aim is to ‘promote uniformity and fixity in the naming of agricultural, sylvicultural and horticultural cultivars (varieties)’. The term culton (plural culta) is also mooted as an equivalent of the botanical term taxon. The Cultivated Code governs the names of all plants which retain their distinctive characters, or combination of distinctive characters, when reproduced sexually (by seed), or vegetatively in cultivation. Because the Code does not have legal status, the commercial interests of plant breeders are guarded by the Council of the International [46]

THE NAMES OF PLANTS The Names of Plants is a handy, two-part reference book for the botanist and amateur gardener. The ...

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