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An Illustrate) ^ Flowering Plant Morphology ADRIAN D. BELL With line drawings by ALAN BRYAN s>< Plant Form An Illustrated Guide to Flowering Plant Morphology Noranteaguyanensis Apitchershaped leaf (bract62) isassociatedwitheach flower; see Figs 88a, bforearlydevelopment. An Plant Form Illustrated Guide to Flowering Plant Morphology Adrian D. Bell School ofBiological Sciences University College ofNorth Wales With line drawings by *m&m?m#m Alan Bryan S0050982 Oxford New York Tokyo OXFORD UNIVERSITY PRESS 1991 . Oxford UniversityPress, WaltonStreet,Oxford0X26DP Oxford NewYork Toronto Delhi Bombay Calcutta Madras Karachi PetalingJaya Singapore HongKong Tokyo Nairobi DaresSalaam CapeTown Melbourne Auckland andassociatedcompaniesin Berlin Ibadan OxfordisatrademarkofOxford UniversityPress Publishedin the UnitedStates by Oxford UniversityPress,NewYork © AdrianD. BellandAlan Bryan, 1991 Allrightsreserved. Nopartofthispublicationmaybe reproduced,storedinaretrievalsystem,ortransmitted,inany formorbyanymeans,electronic,mechanical,photocopying, recording,orotherwise, withoutthepriorpermissionofOxford UniversityPress Thisbookissoldsubject tothecondition thatitshallnot,by Wayoftradeorotherwise,belent,re-sold,hiredout,or otherwisecirculatedwithoutthepublisher'spriorconsentin anyformofbindingorcoverotherthan thatin whichitis publishedandwithoutasimilarconditionincludingthis condition beingimposedon thesubsequentpurchaser British LibraryCataloguinginPublicationData Bell,AdrianD. Plantform:an illustratedguidetofloweringplantmorphology. 1. Floweringplants. Morphology 1. Title 582.13041 ISBN0-19-854279-8 ISBN0-19-854219-4pbk LibraryofCongressCatalogingin PublicationData Bell,AdrianD. Plantform:an illustratedguidetofloweringplant morphology/ Adrian D. Bell:—withlinedrawingsbyAlan—Bryan. 1. Angiosperms—Morphology.—2. Botany Morphol—ogy. 3. Angiospe—rms Morphology Atlases. 4. Botany Morphology Atlases 1. Brijan,Alan. 11. Title. QK641.B45 1990 582.13'044—dc20 90-34783 ISBN0-19-854279-8 ISBN0-19-854219-4 (pbk.) PhotosetbyCotswoldTypesettingLtd,Gloucester PrintedinSingaporeby Times PrintersLtd 'The study ofthe externalfeatures ofplants is in danger ofbeing too much overshadowed by that ofthe internalfeatures. The student, when placed before the bewildering variety offorms does not know where to begin or what to do to acquire information about the plants'. willis (1897) 'Horticulture is, undoubtedly, a great medium ofcivilization, and its pursuit is highly commendable,for it is impossiblefor anyone to study, evenfor a short period only, the structure,forms, and colours ofplants, and benefits derivedfrom the vegetable creation, without an elevation of thought, a refinement oftaste, and an increased love ofnature'. B. S. WILLIAMS (1868) fl have bought me a hawk and a hood, and bells and all, and lack nothing but a book to keep it by'. ben jonson (1598) Coiyphautan Thesinglemonopodialaxis (section250)finallyterminating inaninflorescenceafter44yearsofgrowth. Modelof Holttum (Fig.291c). : Flowering plants exhibit a fascinating array of knowhow, heavily biased towards floral external structures which can be studied with the structure, and has at his disposal a profusion of naked eye or at most a simple hand lens. This is terminology that is daunting to the beginner and Preface the science ofplant morphology, the term being expert alike. A guide is thus required for the used here in the sense that excludes plant benefit ofboth. This book is deliberately, I hope, anatomy. Although an understanding ofthe form attractive, the better to woo the budding botanist and external components ofa plant should be the and the curious amateur plantsman. It is divided foundation ofany botanical investigation, it is into two parts. The first part illustrates and customary to rush ahead, delving deep into the explains much ofthe purely descriptive plant, and thus either ignoring or missing the terminology involved in plant morphology, whilst very features that the plant presents to the the second part deals with an equally important environment. The situation is very well-expressed but largely ignored aspect ofmorphology by my namesake, Professor P. R. Bell (1985): 'In constructional organization. The plant is recent years the spectacular advances in developing, its organs are developing, most molecular biology have generated such flowering plants branch, the branching patterns excitement that there has perhaps been a ofthe plant develop over time, and growth is tendency for organisms to be overlooked. Biology dynamic. Cover ofthis aspect ofplant must nevertheless remain "organismic", and the morphology, which is ofrelevance to the researcherwho loses the concept oforganisms ecologist and the population biologist (Harper seriously weakens his claim to be a biologist'. A 1980), culminates in an example drawn from the blinkered attitude to plants probably commences contemporary morphological world, that ofthe at school level and continues through university. dynamic architecture oftropical trees. The Excellent texts ofplant morphology do exist, but author's fascination with plant morphology has they tend to presume a foundation ofbotanical been fostered by a providential succession of education that is no longer available. The ground mentors, A. D. Prince at school, N. Woodhead at rules ofplant morphology are, by and large, college, and P. B. Tomlinson ever since. Their forgotten (Kaplan 1973a). The^student ofbotany teachings have one principle: ifthe morphology feels this defect but does not know how to resolve ofa plant surprises you, then this is more likely it; the academic conceals his ignorance. It is to reflectyour ignorance rather than an tempting to suggest that many an enthusiastic abnormality on the part ofthe plant. An amateur horticulturist understands plants more unfortunate preoccupation with European plants intimately in terms oftheir morphology than in the past led morphologists to be taken aback does the average botanist. This criticism cannot by the exuberance ofthe world's vegetation, be levelled, however, at the taxonomist who is especially that ofthe tropics. But this is where armed with a great deal ofmorphological the range ofplant form can best be appreciated. mm For this reason the plants illustrated in this book 35 lens) together with extension tubes where Harvard University. These good people have been originate in all continents, and many will be appropriate. Very frequently supplementary light able to point me in a better direction on a unfamiliar to a reader confined to one was supplied by means ofa pair ofsynchronized number ofissues. Any errors that remain are my geographical region. However, the same flash units mounted 15 cm to either side ofthe own; it is inevitable that some morphologists morphological features and details of lens on a bar fixed to the camera body. somewhere will take me to task on points of constructional organization are repeated time Kodachrome 64 ASA was used throughout. I detail, rash generalizations, or personal opinions. and again in totally unrelated plants and the must thank many botanical gardens for allowing I have relied on other people's plant identification reader will recognize familiar forms ifnot familiar access to specimens: The Royal Botanic Garden, in most instances and have followed the names. Edinburgh; The Botanic and Genetic Gardens of nomenclature ofWillis (1973). Let me hide once In a sense this book can be treated as an Oxford University; the Fairchild Tropical Garden, again behind the axiom ofmy teacher at illustrated dictionary to be consulted as necessary Miami, USA; The Botanic Gardens ofMontpellier, university; 'It is the plant that is always right'. and in any sequence. With this in mind, the text France; the private gardens ofM. Marnier and illustrations are extensively cross-referenced Lapostolle, St. Jean Cap Ferrat, France; and the and the index annotated. The seasoned Treborth Botanic Gardens ofthe University morphologist may be surprised to find equal College ofNorth Wales, Bangor. Other space (one double-paged spread) allocated to such photographic sorties have been made hither and an insignificant feature as a stipel (58), and to thither and the author is grateful to those who such a vast topic as floral morphology (146) or have helped him visit various countries in Europe the morphology offruit and seed dispersal (160). and particularly south and central America. I am Whole books have been devoted to these wider very grateful to a great many people who have topics to which references are given, where helped me in different ways to complete this appropriate, rather than the information being guide: Nerys Owen for typing and most efficiently duplicated here. All the line drawings and converting recorded tapes into word processor diagrams are the work ofAlan Bryan to whom I format without complaint andJosie Rodgers for am clearly indebted. Alan's talent as an artist taming the index with a suitable computer represents a happy combination ofnatural program; my colleagues at the School ofPlant ability, an eye for detail, and a classical botanical Biology, Bangor, for their encouragement and in training. Practically all the drawings and all the particular ProfessorJ. L. Harper for recognizing photographs have been taken from living plants, that plant morphology is a key subject. A the exceptions being a few dried woody number ofkind people have commented on drafts specimens, and a very few that have been ofthe manuscript at various stages. I am adapted from existing illustrations. All the indebted to Professor F. Halle ofthe Botanical photographs were taken by the author (except 7 Institute, U.S.T.L. Montpellier, to ProfessorP. as noted) using an old Pentax Spotmatic II Greig-Smith and Dr. and Mrs. N. Runham of mm camera with a 105 lens (or occasionally a Bangor, and to Professor P. B. Tomlinson of