MANUAL OF HERBARIUM TAXONOMY THEORY AND PRACTIC 2 Edited by: E. F. DE VOGEL Rijksherbarium Leiden, The Netherlands UnEsCO United Nations Educational, Scientific and Cultural Organization Regional Office for Science and Technology For Southeast Asia Jalan M. H. Thamrin 14 Jakarta 10240 Indonesia 1987CONTENTS PARPACE Cee peas: ACMAONLRLORUSWTS Stunna INTRODUCTION. 0 ee 1. ‘THE TWO CHALLENGES FOR PLANT SYSTEMATICS (Clkatkeany co uenci a 1, COLLECTING (M.M.J. van Halqony) 0 TET, MANUAL FOR THE DESCRIPTION OF FLOWERING PLANTS (J.F, Voldkamp) . . . . wee ee Tila, SEQUENCE OF ORCANS AND TERMINOLOGY OF CHARACTERS (J.P. Voldkamp) . . . . PES TItb. INDEX TO TERMINOLOGY ().F. Voldkagp) . eee IIc. ABBREVIATIONS USED IN ENGLER & PRANTL (J.P, Veldkamp) . . Tild. EXAMPLE OF AN ANALYTICAL DERCAIPTION (B.F, de Vogel) Tile, GUIDELINES FOR THE PREPARATION oF REVISTONS 1V. STRUCTURE OF KEYS FOR IDENTIFICATION (R.Geosink). v THRORY OF CLASSIFICATION OF ORGANISMS (R.Gvostnk) VI. BOTANICAL NOMENCLATURE OF YARCULAR (LANTS (J.P. Voldkasp) . VIL, ADMINISTRATION Vila. THE PAPER FOUNDATION (J.P Velidkamp) VETb. CITATION OF MATERIAL (J.F. Veldkaap) Vile. COLLECTIONS CITED UNDER ABBREVIATIONS OL dacobay 20 26 65 1 "A mw m a 127 447 155 158‘The authors of-the papers published In this volume are solely responsible for the choice and presentation of the facts presented and for the opinions expressed therein, These are not necessarily those of UNESCO and do not commit the Organi- zation in any way. ‘The designations employed and tho presentation of material throughout the publication do not imply the expression of any opinion whatsoever on the pat of UNESCO concerning the ivgal status of any country, territory, city or area, or of its authoritles, or concerning the delimitation of its frontiers or boundaries. 1PREFACE In September 1985 & Training Course on Research Methodology in Plant Texonomy was conducted in the Herbarium Bogoriense under the auspices of the former National Biological Institute - LIPI, the Rijksherbarium Leiden, and UNESCO-MAB. The course wes attended by Indonesian junior sclentists and junior lecturers at the beginning of their career and specialization as practicing plant taxonomists. ‘or that purpose 5; hand outs were prepared and accumt id into indy manual which was consulted daily by the trainces throughout the course, From the very beginning it was felt that the’ book will ‘be useful for others venturing into this branch of botany which nowadays is neglected if act altogethsr dered outmoded, Since UNESCO and the Indonesian National MAB Programme are Interested in promoting plant taxonomy in Southeast Asia, as evident from the fact that both organizations will cosponsor a rather regular "Regional Training Course on Plant Taxonomy and Botanical Exploration towards the Preparation of the Flora of Biosphere Reserves in Southeast Asia", it is decided to publish the manual to make it available for a wider audience, Dr. EF, de Vogel of the Rijksherbarium Leiden voluntered to edit the Manual as’ it Js presented here to the public, The interest of UNESCO in this field has been of long standing because as early as 1960 this agency already conducted a similar training course in Ujung Kulon (West Java), and about half of those attended that course nowadays are still active as field botanists and plant taxonomists. It is hoped that the publication of the present Manual will attract more potential students in this field so that in the future plant taxonomists will not always remain a scarce commodity in the region, Mien A. Rifai, Pustitbang Biologi-LIPI, Bogor Kuswata Kartawinata, UNESCO/ROSTSEA, JakartaACKNOWLEDGEMENTS The following akncowledgements are gratefully made for texts and figures: Flora Malesiana (text on p. 79); Blumea (texts on pp. 80, 84 and 85); Dr. Bertel Hansen (figure on p. 80); and Mr. R.J. Johns (figure on p. 81). VILINTRODUCTION ‘This manual wee originally drawn up in 1985 as a written aid to a course in Plant Taxonomy for a rather heterogeneous group of par- ticipants, consisting of young staff-members of the Herbarium Bogoriensa and younger lecturers of some universities and biological institutions in Indonesia. We made the manual with our own advanced students in mind, which means that it contains the sort of knowledge we like the students to possess when they prepare a revision of a group of cies for the Flora halosiana. This ‘poi ion of knowledge’ should not be taken too literary: it is NOT necessary to learn all terms and their meaning(s) of chapter III by heart). it 18 neceseary, however, that you know where to find them (via the index), and what sort of terms are explained where. Similarly, while preparing a part of a flora or a floristic survey, it is NOT necessary to know exactly, how to make cladistic analy: but we think it IS necessary that you understand the difference between keys and classifications. We algo think it necessary to realize that different kinds (phenetic and phylogenetic) of classifications represent different aspects of the knowledge of Nature (similarity-classes or nested sets of individvals) on the levela of Organisms, Species, and Higher Natural Taxa. : Chapter I is an overview of the aims of systematics. One challenge is the question: What is the Natural Classification uf the group under atudy ?! The other challenge is the question: ‘What is the course of (phylogenetic) history that lead to this Natural Classification ?! This chapter is meant as a perpetual question (yes, one question indeed): What am I doing as a taxonomist ? Don't read it once, read it over again, and you'll read different matters in it as your experience grows. Chapter II is lese theoretical) in it is described how modern, useful collections should be made, It is useless to collect a small fragment of a common wayside tree from W-Java, and add it. to the collection of any scientific Herbarium. But if you collect a rich specimen, with flowers and fruite, male and female flowers both collected, also in PAA, with colourslides and provided with relevant field-notes, well, + « + SUCH collections are welcome, even of very common species. Chapter III is also mainly practical. Terminology is not a matter of Nature iteel£, it is rather the set of agreements among professional taxonomists so that scientific communication is possible. The applied terminology is different per plant group. Example: grasses have paleas and lemmas, not sepale and petals, Grassee do have organa that are perhaps homologous to aepala and petals, but they are not the puleas and lemmas, and homology ia a theoretical maitar, Terminology is learned by practice, but simultaneously theory-dependent. We advise to exercise terminology’ by making analytical and diagnostic descriptions of a few differently structured plants. Chapters IV and V are mainly theoretical, Like in the case of chapter I, we advise not to go through them just once, Read and read them again, and, more importantly, read (and reread) the selected litera~ ture enumerated at the end of the chapters.Chapter VI deals with the mainly ‘juridical’ matter of Nomenclature, The aim is to have relative stability of scientific names: One accepted taxonomic classification laads to only une correct name per accepted taxon. The internationally accepted principles are priority and the type-method, and in this chapter it is explained how to apply the 75 articles of the International Code of Botanical Nomenclature, based on these principles. Chapter VII contains the basic administration necessary for the preparation of a classical revision in tl of a regional Flora-project, Minimally such an administration consists of lists of studied specimens, lists of literature references and above all, a practical arrangement of your own observatione, The presentation is based on the gond-old paperindex system, of the heterogeneity Of computerized systems in use today we have refrained from presenting administration in a rore modern way. Remaine the question how to apply this manual, We use it as an aid to a course roughly consisting of three parte: 1, First get acquainted with terminology of the parts of ‘normally constructed’ flowering plante (no gras palms, or Rafflesias, but instead plants with usual leaves and bi 1 flowers with obvious calyx and corolla) by means of constructing analytical descriptions of not too complicated flowering plants, This part is gradually connected to the secund part by means of the first exercises with the iden- tification of the described plants. For ‘these purposes we use "thonner'a key to the families’ ,a regional or local flora, and chap- ter III of the Manual. 2. The integrated block, This part of the course deals with the question: 'What things can you do with character-analyses ?' Firstly, eight ready-made diagnostic descriptions are identified by means of keya. Then some more plants are described (from dried material) in a similar way, and comparable character states are put in a datamatrix, From thie datamatrixa: constructed: a key for identification, a phenetic classification, and an attempt to the natural classification of these few plante only. The chapters IV and V of the manual are read in this part of the course. 3. The mini-ravisions, This part of the course deals with the delimitation of species, infraspecific variation, and perhaps (infra-) generic problems (depending on the subject). An amount of dry her~ barium material is presented with the questions: how many species are represented by this pile, how to distinguish them, what are their proper names, and what is the possible relation between the species ? During this part of the course the participants get acquainted with most aspects of a classical revision, the base of regional floristic work. For this purpose, the remaining chapters (1, VI, and VII) are atudied, and we hope, that during the practical work, the relation of all chapters will have become clear to'the participants, Plant Geography is lacking in this manual, due to the limited amount of time available for the preparation of thie manual. We hope to provide a chapter on this subject in a next edition.I. THE TWO CHALLENGES FOR PLANT SYSTEMATICS (Cc, Kalkman) RELATIONSHIP The central concept in plant systematics (as in the systematics of other organisms: animals, fungi, prokaryotes) is that of relation~ ship, Different kinds of plant taxonomists, engaged in differant kinds of research, will subscribe to the statement that searching for rela- tionships is an essential part of their activities. Plant systematics can, therefore, rightly be d-¢ined as tha science of relationships be- tween plants - or rather: between plant populations. There are relationships between populations living in the same time, and between populations in different times, horizontal relation- ships and verticel ones, See the top of figure 1a, where tha “horizon- tal" relationship is -indicated between the two populations Cc and D, both Living in one time-level: at present. 4] Howe == relationship 3 b a course of evolution Pig, 1, Relationships and genealogic connections. In a the two- sided arrows indicate the relationship existing between the popula- tions, In b the one-sided arrows chow that these relationships can have different historical backgrounds. See also fig. 4, 1If population C is (fig. 1) closely related to D this implies a number of things. From such statement one can draw the following conclusions: 1 that their genomes (genetic composition) will be largely identical; 2. that individuals from C vill be able to hybridize with indivi- . duals from Dy 3. that C and D had one coemon ancestra) wopulation, not too long agor 4. that the individual plants from C and D will be much alikey Be that C and D must belong to one taxon of a low rank or to two taxa of low rank which are placed closely together in the classification. In the same figure 1a the populations A and B have been drawn as living in the past (see time scale on the left), The %wo-sided arrows in this figure do represent genetic relationship, nc_ the course of evolution. This becomes obvious in comparing figure 1b where the ar- rows axe one-sided and are meant to indicate the course of the evolu- tion. Then we see that two different evolutionary trees can be repr. sented by the same diagram of relationships. Let us return to fig.» la, If population A ia called closely related to population B (living in a different time-level), this again means a number of things: 1. that their genomes will have been largely identical (although we will not be able to prove this genetically), 2. that individuals from A and B could have hybridizod if they had lived next to one another in the same time (and also this cannot be yroved)) 3, that there is a genealogical tie (a familie-tie) between A and 4. that A and B wil be much alike; 5. that A and B must be classified in one species. The two lists are obviously alike. We will consider the points nore closely. Point 4 means that the term and idea relationship can be used also if we know nothing about history but only have the differences and similarities at our disposal. Tha notion dates back from before the development of genetice and evolutionary biology. In what follows, we will not use the term in that pre-evolution way, i.e. not in the sense of “similarity” only. Points 1 and 2 indicate that the idea relationship nowadays has 4 distinet genetical load, aleo when it has not (yet) been posaible (oc will never be possible) to find out the actual genetical situ tion. In point 3 evolution is introduced as determining the degree of relationship. We will return to thie point later. Genealogical connec~ tion (connection by descent) is what we imply in everyday life by using the word relative, more or less the same is implied when using the word relationship in systematics, CLASSIFICATION AND PHYLOGENY The points 1 to 4 are not independent from each other: they aredifferent aspects of one phenomenon, Evolution, genome, likeness are avidently linked to each other, Point 4, however, introduces quite a different basic consept: classification, Classification is the distinguishing and naming of unite (taxa) in a hiezarchicei system, usually simply called "the" system, The part of systematic, particularly involved with classification, one can dif- ferentiate as taxonomy, but often the terms systematics and taxonomy, aystematist and taxonomist are seen and used as synonyms. Taxa have different ranks in the system, from subforma to reg- num, the species is the conceptually most important category, Apart species, only occurring in one place, an extant species number of populations that can be more or less distinc- tive. When Gistribution ia spot-wise, as in a mountain area or in an archipelago the populations are sharply. separated and may show differences; when distribution is more continuous, the populations are often not separated and usually grade into the neighbouring ones. A species has spatially a certain extent, but also in time there im continuity. Thie is symbolized in figure 2, where is shown that Populations of one species on one time-level can be different (symbolized in divarse kinds of shading). This variation undoubtedly has, among others, to do with ecological factors. There are also changes in time, as was already treated in the discussion of figure 1. ‘the history of changes in time is called phylogeny, Fig. 2. A species in the course of time, Different shadings indicate thg& the populations are .not quite identical and together define the Wiriotional amplitude of the species. Also is seen that populations may appear and disappear in time. Expressed in terms of populations, in the course of time popula~ tions of one species can become separated. The cohesion of the an- cestral populations has diminished and the later population cannot any longer be placed in one speci: The causes of these differences can ber: mutation, hybridization and polyploidization, in combination usually with geographical or ecological isolation and changes in the environment giving differences in selection pressure. The study of these mechanisms ia the subject. of population genetics and is not elaborated here In figure 3 is visualized how species originate out of on ai 3cestral species (speciation). The situation as drawn can be “translated” in different wayo in the classification, see the legend, a @ at first b later Fig. 3. Speciation, KLMN is a species with itn popilations, 8 is another, related species. At a later time the populations K and M have diverged, K' and M' are more different than K and M originally were. One may decide that K' and M' are indwpendent species now, or subspe- » Population L is extinct, Population N has drastically changed by hybridization (the black arrow) with a population of species 8, which was followed by polyp’oidization, That, brought species (or subspe~ cies?) P into existence. : It will be clear now that also on a level higher than the population one can speak about relationships. Two species may be called closely related, ift 1, dt can be supposed (or it has been shown) that the genomes are more similar to each other than to any more remotely related species; 2. hybridization (exchange of genes) batween ind. viduals of the two species is still possible or at least not abzoiutely impossible, although maybe only after technical tricks or although the off- spring may have lesa vitality or fertility; 3e it maybe supposed that the species have a common ancestor. 4. The close relationship is inferred from the observation that the individuals of both species are much alike. Higher up in the system, speakihg about genera and families, the term rolationship may still be used and be useful, although the list of criteria becomes shorter and more vaque. In taxa above the speci only supposed genealogy (point 3) remain, inferred from likenese (point 4).THE TWO CHALLENGES Now we can start elaborating the two challenges that systematics and the taxonomists have to face, Problema in systematics can always be placed under ona of two headings, however different the practical ‘research goals may be, They are the following: 1. ‘The phylogenetic challenge, The question asked is: How can one imagine the populations/taxa un: vestigation to be intercon- nected in time? This of course depends on (biological and other) phenomena having their influence on the course of evolution, These phenomena can be geographical isolation, change of clima- te, ecological differentiation, hybridization, polyploidization, eto., as already indicated above, An opinion on relationships depends in this context on knowledge of or hypotheses on the e- volutien of the group and the biotic or non-biotic procesnes in- volved. 2, ‘The olassificatory challenge. The question asked ie: what is the beat classification of the taxa involved? This is not a purely technical-administrative question, it is not just an art or hese are certainly nt (see In many cases the claseification will entirely or partly, explicitly or not, be based on knowledge of or hypo- theses on phylogenetics, as mentioned under 1, Sone taxonomiate, however, try to heep phylogenetica out of their classification, although this is very difficult. The two kinds of questions asked are obviously not identical, although in the daily practice of actual research they are oftun found to be hardly separable, CHARACTER ANALYSES ‘The data which the taxonomist needs to guide him to an answer on hie questions, have to be gathered from: - study of living individuals, either in the field as parts of natural population(s) or in garden, greenhouse or culture room as samples from natural populations, and/or from: : > study of conserved individuals, sometimes fossils, usually herbarium specimens, i.e, omall and dead samples from populations that the ro~ searcher d'd or did not study himself. ‘The nature of the data to be interpreted is varied and so are the methods used in collecting the ® Visible characters can be studied by anatomical techniques. "Vieib: chromatogram. can make visible an inviaible compound, microscopa has made visible what a short time ago was invisible. If we restrict ourselves to the naked eye, the hand lens, the light micro- scope and the (scanning or transmission) electron microscope, then structural characters can be studied of organ systems (e.g. flowers), organs (e.g. sporangia), tistues (e.g. xylem), cells (e.g. spores or unicellular organisms), cell organelles (e.g, chloroplasts, chromoso- mes). Systematic anatomy and karyosystematics (cytotaxunomy) are spe- cialist approaches on thin level.Non-visible characters can be detected in the chemical processes (and the resulting products) in the plant, but also in their "physiology", i.e. the inbuilt responve to environmental factors, de- termining ecological preference, reproductior. etc, Generally the sys~ tematist will he more interested in the products of physiological and chemical processes than in the processes themselves ~ even although it ean certainly defended that the latter are the real characters ince the outcomes (e.g. accumulation of a certain compound or sporulation at a certain temperature) can be produced in different waya, A large part of chemosystematics is concerned with the distribu- tion of secondary compounds like e.g. essential oils or alkaloids, The molecular structure and composition of the chromosomes may - theareti- cally - be considered the most important criterion for relationships: all processes and products are derived from them, Slightly less fun- damental in the life of the plants are those enzymes of which che for- mation is regulated directly by the genome, As far as morphological/anatomical and chemical characters are concerned, there is for many plant groups alrearly much insight in the degree to which character state: genotypically dotermined and which ones belong to phenotypic variability, Referring back to the earlier lists: it can often be stated with confidence which likenesses (point 4) in morphological or microscopical structures or ir chemical composition may be interpreted as a result of genetical identity (point 1). In systematic studies with living plants ification is possible by means of cultivation under controlled circumstances, Reproductive barriers (point 2) can of course only experimen- tally be proved to exist or not and that brings us from descriptive to experimental techniques. I use the term experiment in the sensa oft interventions of the scientist in the natural course of affairs, with registration of the consequences, in order to prove or make plausible a certain causal relation, The only experiments that the taxonomist can do are either ecological (tcansplantation, cultivation in a con- trolled environment) or genetical (hybridization efforts). Experimen- tal systematics wav originally aptly called genecology (Turesson 1920- 30). As in most disciplines atatistical handling of data has become very important. The application of statistics to a multitude of data ig in itself not a new development but computers have made practicable much more than before, We have, however, than switched from the re- trieval of data, to the processing of them. The entire field indicated above, be summarized as the character analysis, This analysis, mre or less complete or very one- aided, is available for finding an answer to the question(s) posed. So we return to the two questions, the two challenges, the two commis- sions of systematics WHAT IS THE BEST CLASSIPICATION ? In the first place: what is the beat way to arrange the taxa in @ aystem of classificction? ‘The structure of the system is’ given: it is a hierarchy of the categories forme, variety, subspecies, species, genus, etc. and (ac~ cording to the international rules of nomenclature, art, 2.1) every plant “is treated ae belonging to a number of taxa of consecutively subordinate ranks, among which the rank of species is basal". Apart from this fornal (taxonomic) system there exivt also in-formal systems where the rules of nomenclature are rot valid and which do not have to be hierarchical either, With Danser (1929) we can dis- tinguish the entities conmmiscuum and convivium (gunetically defined) or following Gilmour & Gregor (Nature 144, 1939: 333-334) the cate- gories cytod-me, ecodeme, gamodeme, etc. (genetically and ecologically defined). These categories demand knowledgu on the level of population and are only employable when living, wild populations have been stu- died, ‘There ie a number of criteria to which an “ideal” classification should answer: 1 Tha taxa must be identifiable, i.e. they have to be distinguieh- able also for.others than the classifier, 2. ‘The predictiva value must be high, in the sense that as yet unknown specimens/populations must fit into the classification. 3, The biological value must be high, in the sense that the class ification must “be in accordance with 6.9. ecological prefer- ences, with diecrimination by parasites or predators, and with biological (im)possibilities like crossing behaviour, 4. ‘Thephylogenetical value must be high, in the sense that 411 taxa must be monophyletic (= allparts of the taxon are descendants of one ancestral species) and that the hierarchy mirrors the phy- logenetic history. An ideal classification does not exist, 10 more than other ideal situations, Also a taxonomist can do no move than strive at an ap- proximation of the ideal at some points at Jeast. What happens during the process of classifying, can be sum- marized quite simply: one studies characters and properties, with the aid of one or more of tho available techniques as mentioned in a pre- vioue paragraph, weigh similarities, differences and the outcome of experiments, and at tne end of the atudy one unites populations (or individuals} to species that must be placed in a genus, a femily, ete., and/or that car. be divided again into varieties, etc. The classification rests on a firmer foundation and can become More robust, when move characters are involved. However, this does certainly not imply that decisions are easier then, Sometimes parts of the character analysis iio not (seem to) conform, The weighing process becomes more difficult then and often a solution is expected from a broader factual basis Angluding a different Kind of characters, in the hope that the causes wf the “conflicting evidence” will become clear and that the conflict will disappear. The units one ‘aa to work with, the taxa from low to high, may be fixed, about the contents of these concepts a number of questions can be ed. Partly the: questions are philosophical (does a species, a genus reilly exist in nature or only in the human mind?), partly they are quite practical (to which criteria must species, genera etc. answer?). Especially about the species’concept there has been lot of discussion and opinions did and do differ. For many a species ig a unit that has to be morphologically recognizable with preferentially two constant differences with other species, for others the species ia more or leas identical with the hybridization com- munity. Genetical "identity" of the components of a species is an im- portant criterium and may lead some to forbid different chromouome numbers within one epecies. The battle about "biological" and “taxonomical for less clear even: linnean)" species started in the forties wien herbariam taxonomists were confronted with aetudents of 7living populations and when a "new systematics" entered tha field with ite emphasia on what happens with the living plants in the field. The fire of the battle has subsided now and a more synthesizing movement ham replaced polarization. ‘The question was: what is the best classification? That suggests either an objective standard outside the classification ayainst which the latter can be tested or a tolerated subjectivity that also to- lerates that not all scientists necessarily must come to the same con- clusion, Despite claims heard from time to time that “here are methods to read from the objective data of character states the classification in an objective way, in my opinion it is the latter, Numerical taxonomy has developed methods to express numerically the similarities and differences between individuals, populations or clusters of them, As long as the same character states are scored in the same way and clustered according to the same rules and if it ic agreed at which number or formula a grouping may be celled species, variety, etc., a classification is repeatable indeed. That it is then much more objective than non-numerical methods, is questioned by many. A subjective evaluation (a.cording to tradition or revo- lutionary) of the characters, of their similarities and differences, by the scientist himself, seems to be unavoidable, It is this subjec- tivity that makes classifying a bit like an art (the term used as in “arte and sciences"), Different people must, if they measure with dif- ferent methods the specific gravity of mercury, the molecular weight of sulphuric acid or the refraction index of benzene, come to identi- cal conclusions but different people may defend quite different sys- tema - just as students of literature or history do not necessarily draw the same conclusions when they look at the same facts. There is, however, no reason to be too sombre about this. In well-known plant groupe like the Seed Plants, there has developed a distinct common opinion about what can be distinguished ao species, genera, families, The differences in opinion are, as always, more con- spicuous than the unanimity: more ia written - to mentione one sample - sbout the question whether the Yellow Archangel belongs in the gonus La.ium or in a separate genus Lamiastrum than about the common opinion that the c. 25 other Dead-nettles are species of Lamium. In less well- known groups, G.g. Chrysophyceae, texonominte of different generations or in the sam- period can certainly come to different views. These differen: @ then caused at least as much by a deficient knowladge of necessary fact than by the subjective handling of the facts. “Lumper" and “splitter” are the predicates used by the taxonomists for their colleagues who in their opinion give the wrong enphi to differences resp. similarities. This kind of derogative terminology can better be evaded. WHAT IS IE BEST PHYLOGENY 7 In what way can the phylogenetic history of the group best be reconstructed? This question can ba asked for its own scientific value but also in order to use the answer in the classification (see p. 7 under nr. 4). Phylogenetical research can be done on different taxonomical levela and with different means, The factual data on which to base the phylogenetical hypothesis are the same as those emerging from the character analysis that is used in classification. On the lowermost taxonomical level, that of the species, the in- fraepecific taxa and the species complex, the micro-evolution can be studied. This is the domain of biosyatematics, where experiments arecombined with karyological observations (the chromonomes: nunber, shape and dimensions, behaviour during meiosis), The results of population-ganetical reserach end more in general of evolutionary biology (atuay of the processes leading to evolution = changa in time, eepectally the study of apeciation) are foremost in biosystematics, On a higher taxonomical level, that of genera and higher categories, the disciplina studying evolutionary change is called phylogenetics. The ground is less firm here because experiments cannot be done, relation to popuiation ee je indirect, the time period in which the evolutionary processes took place is longer and longer ago and this all gives more room for speculation. The last word has to be understood as: the framing of a hypothesis that as well as possible fits tl known data, that does not ask for unknown evolutionary proc and that 4s supporind by data from palaeogeography, Fossils may belong to the "known data” but problems with reconstruction of fossil plants, with dating them and generally the scarcity of fossils make the help of them-often not more than marginal. Reconstruction of group phylogeny is realized, if not quite in- tuitively, by using the most probable character phyloyenies (tran: formation series) and this makes for big problems. Not even knowing yet through which molecular and cellular processes organa are formed and transformed during ontogenetical development in the present (morphogenesis), how is one to make sure how the organs have been transformed in the past (phylogenesis)? Even obvious statements like “free parts are preceding connate ones" will doubtless have their ex- ceptions, i.e, raversals, On the possibility of giving a sound judge- ment on the problem of primitive/derived, general/specialized, plesio- morph/apomorph (or what terms one may prefer) different people think very differsatly, 1.e. from optimistic to pessimistic. Cladistic analysis according to Hennig and his followers is the most formalized one among phylogenetic reconstruction methods. It fol- lows strict rules but also in this method the truth does not come as a matter of fact: making choices cannot avoided. However, it is a very clear method where the choices are explicit, not hidden away and therefore open for discussion and dissent. In phylogenetics circular reasoning (also auphemistically called mntual enlightenment or positive feedback) is hard to evade, even when one tries explicitly to bridle ones "intuition". As long as the phylogenetioist realies that absolute verification of unique happen- ings in the past ie impossible, as long as he does not think that through adding up three or four plausibilities or possibilities cer- tainty emerges, as long - consequently - as he realizes that a phylo- geny has the character of a scientific hypothesis which can be tested, corroborated or verified, there ig nothing wrong with phylogeny. Toeting a phyloganetical hyjothesis can (except sometimes in biosyatematics) not be done by experiment but muat be done by further observation, ¢@.g. of other character complexes or using other tech- niques. The difference between experiment and observation is, however, not as fundamental seems to be at first sight. Thus it 1s possible to find an anawer in the search for the best reconstruction of the phylogeny in a certain case, a preliminary anewer, it is true. But does science ever give final anawers? ‘A phylogenetical hypothesis can be visualized in different ways. The most exact one is the phylogenetical tree, with the time as one of the axes. The tree is usually placed vertically with “now* at the top, Fig. 3b and 4a are examples. Mot all drawings locking like trees, however, have the time 9an axis and do not always picture "descent", They may also be a fun- damentally different aymbolization of the degree of similarity, We saw before that similarity and phylogeny are not independent of each other but they are certainly not identical concepts, Seeing a tree-like sym- bol one must, therefore, always try to realize what exactly has been pictured (seo legend of fig. 4, where also some other symbolic draw- ings are presented). + abd c d ~ taxa living now abc d- taxa V " 260 © on Fig. 4. Phylogenetical trees and relationship schemes. In a a tree of four taxa has been drawn, a further simplification of a figure like 3b. There is a time axia and the tree symbolizes the historical events as the author thinks they have been. In b and ¢ relationships are drawn without a time axis, In b the distance between the circles or rectangles indicate the degree of relationship. Connecting lines, interrogation marks, etc. may be used to accentuate the situation as hypothesized, Between contemporaneous groups no arrows must be drawn, A scheme like ¢ (dendrogram) is very much like a phylogenetical tree but lacks the time-axis. Dendrograma ara drawn e.g. in aumerical taxonomy and the length of the connecting lines indicates the si- milarity between the taxa: a and b are very similar, the likeness with o ia lesa, etc. Introduction of a time axis gives such a dendrogram a Phylogenetical load. 10THE PRODUCTS Publications in plant systematics sometimes are built according to the stereotypical pattern in natural science: definition of tne problem - material and methods - results of observations and/or ex- periments - discussion. In many cai however, the results of the systematic research are presented differently, viz. as a survey of a part of the system of classification, The degree of elaboration of the aurvey and the scope of the research (geographically and taxonomical~ ly) may differ widely. The diagram in fig.°5 gives a diagram of the different types of classifying publications. eraphi | WORLD + REGIONAL —————-—-»_ LOCAL monograph floristic revision Fig. 5. Publications in plant systematics. “Regional” are publi- cations on areas like Europe, tropical Asia, Australia. “Local” means a coverage of e.g, the Netherlands, Java, «ifornia, Nain groups are Algae, Mosses, Vascular plants, Angiosperms, etc, Small groups are or- ders, families, genera. Transitions between the types of publication are not sharp. Variation in depth is determined by a) the degree to which dif- ferent techniques have been applied in: the character analysis and (related) the kind of characters involved, and b) the degree to which the basal character analysis is made explicit or left implicit in the descriptions of recognized taxa, In the monographs one expects an ex- plicit, elaborate character analysis and an elaborate discussion of the taxonomical conclusions drawn frov, it. In small floras of course there is no room for this. In monographé also the phylogenetical prob- less are often discussed and the large systematic surveys are nowadays usually strongly coloured by phylogenatical speculation, Apart from the kinde of publications as given in fig, 5, there are also reports on the study of only one character complex, e.g. al- kaloid pattern, vascular system, flagellar structure, We are then at the boundary between systematics and ancther discipline, e.g. organic ilchemistry, plant morphology or cell biology. The degree to which the systematic problems (phylogeny or classification) are playing a role in the actual research project, varies highly from case to case. Pub- Mecations of this type usually follow the stereotypical pattern, THZ NATURE OF SYSTEMATICS In former days a’ difference was made between general botany and special botany pecially under German influeaces: allgemeine va. spezielle Botanik). Special botany covered plant systematics and plant geography,’ whereas morphology, physiolegy and ecology belonged to the general branch. The subdivieion is practically obsolete now but it may exve us to stress a fundamental quality of systematics and historical biogeography, Both discplines are not - as most other subdiaciplines of biology - aiming at the discovery and formulation of general laws but at the explication of special cases. Systematics does not want to elucidate how in general evolutionary changes are taking place (the field of evolutionary biology) but how in the case of genus A, family B or species C evolution did happen. The special casa, the one-time occurrence is what counts. In this light systematics is a historical science. Subject of research ie the pluriformity in the living nature, a pluriformity that originated because certain unique events (migration, change of climate, mutation, hybridization, polyploid! zation) in a certain constellation have led to certain unique results. , With the aid of observations and experiments in other fields (ecology, genetics, evolutionary biology) one can start to apprehend more or less how things happened, Complete certainty, as pursued by physics, cannot be reached because of the irrepeatability of history. What is said here about systematics, is also valid for historical biogeography where knowledge of the earth sciences and ecology may bring into reach the global reconstruction of the history of the areas of species A, genus B or family Cc. A “theory” in these disciplines never runs “it is always so and so” but “it may have happened so and so“ - not the inevitability of the (dead) natural laws but the probabilities of the events in the living nature. THE USE Sometimes people ask: What is the use of plant systematics? The anawer is two-sided and the usefulness of plant systematics has two very different aspects - maybe not always strictly distinguished, In the first place there is the usefulness of taxonomic knowledge: it is aften recognized that knowing planta and their names is an ential aid in many kinda of human activity (from forest exploitation to na- ture conservation, from medicinal application to weed control). Th call for more taxonomists is, therefore, often heard. In the second place the usefulness of taxonomic research and its priority over other kinds of earch can be questioned. People in Western Europe may be led into error since their own flora is rela- tively wall known, So they forget that there are countries in the world without even a Flora for the ‘Vascular, Plants, not to mention other plant groups. Mostly th countries are not well-explored either, There are many genera and families of which the taxonomical knowledge has for the last time been summarized around the year 1900 4nd that can of course only be a very incomplete and inadequate syn- thesis, There are still seaweede of which the life cycle is unknown 12and Angiosperm species of which only the male flowers are known, not: the female ones. Well-founded decisions about relationshipe will of course be difficult or impossible then. Only of a small minority of species (mostly herbaceous plants of Northern temperate regions) the population genetics has been studied, To summarize: in the entire dis: cipline of syatematics comprising the character analysia, the cla iffcetion, the phylogeny, there is still plenty of innovative and im- proving research to do. ‘That reserech will lead to enlarging, broadening and deepening of the available knowledge, not only available for other taxonomists but also for students of other disciplines (biological or not) and for practical and applied purposes, The practical knowledge can only remain adequate and useful, also in future situations, if the research remaina alive, active and modern. EPILOGUE Quite at the beginning of this chapter I mentioned “different kinds of plant taxonomiste engaged in different kinds of research". I hope that indeed it became clear that under the roof of plant sys- tematics there is a large diversity of activities and purpos may find a plant taxonomist at work behind a chemical apparatui tween piles of herbarium specimens, in a garden between flowers en- eased in gauze, in a cold culture room between bottles with seaweeds, in a tropical forest, as well as behind a scanning electron microscope or a computer. T am aware that this short text is only a meagre representation of the richiesses of plant systematics, Much has not been mentioned, stress was placed on the two-sideness of systematics, the two kinds of questions one can ask, the two challenges confronting the taxonomist: ‘The differences have been stressed, I: hope that also the fundamental unity has become visible. This is a translation of “De twee vragen van de plantensyatematiek", _ published by the Rijksherbarium (1982). 13II, COLLECTING (4.M.d. van Balgooy) INTRODUCTION ‘The preparation of a good collection is an essential aspect of herbarium practices, The following lines are intended as a brief in- troduction to collecting. For a more detailed treatment of the sub- ject, consult van Steenis (1950), Fosberg & Sachet (1965) and Womeraley (1976, 1981). ‘A good herbarium specimen must give the investigator the best possible information on the plant concerned, In other words, the col- lection should preferably contain atl parts of the plant and should be well annotated so as to provide all information not visible on the herbarium specimen. HOW TO COLLECT Small plants should be collected as a whole, Larger terrestrial herbs, shrubs, creepe: and small climbers can easily be collected with secateurs or pocket knife. Firmly rooting or cuber bearing plants can be dug out with a pick or a parang, Most of the lower trees can be harvested from the ground with a pruner. Real problema are presented by tall trees, lianas and epiphytes growing in thair crowns, high above ground, The best way by far to collect these is by a tree climber, who can select the best material without undue damage to the plants. It ie often difficult from the floor of the forest to detect whether a tree or liane, is in flower, A good pair of binoculars is indispensable. Often one finds fallen flowers and fruits on the ground but it is not always easy to ascer- tain from which plant they have come. If it ie for some reason’ impossible to climb the tree, other methods must be applied, One of the cheapest methods is by using a slingshot (catapult), A sinker at the end of a nylon fishing line wound around » casting al is shot over the branch to be collected, After removing the einker the fiching line is connected to a stout cord, which is pulled over tt) branch by winding the fishing line back onto the reel. By pulling both ends of the cord the branch can be snapped off, See also Hyland (1972), The branches cf some trees (Myristicaceae, Moracéae) are very tough and will not break. In thie case a wira saw can be inserted in the middle of the cord. The branch can then be sawn off by alternately pulling each of tha two ends of the cord. Shotguns and crossbows are also in use for the same purpose and elaborate olimbilig devices have been applied, but are mostly too ex- pensive and too cumbersome to carry for our purposes, See e.g. Perry (1978), If all other methods fail, the tree has to be chopped down, a practice to be avoided, especially in nature reserves. An advantage of chopping a tree is that it yields ampie material to choose from, Fur- thermore, in its fall the tree often brings down lianas and epiphytes that would otherwise be missed. 14WHAT TO COLLECT Ag a general rvie do not collect sterile material! It is mostly useless for scientific purposes, and identification - if at. all pos- mible - is often very troublesome and time-consuming. deally a collection should contain all parte of the plante: fruits, 1 » wood ete., but a branch with leaves and flowers or fruits is in most cases sufficient for identification. Col- lect a fair-sized sample and not a scrap. Hany plants ace dicecious, collect both male and femal nt. Some speci: , for instance leaves in Juvenile specimens or on sucker shoots may be large and strongly dissected, whereas the adult plant may have small entire leaves; collect both! If a plant ie parasitic try also to collect the host. This may not be easy in root parasites Balanophora, When collecting lianas try to find out their means of climbing: hooks, tendrils, special branches, adhesive roots etc. See also Jacobs (1976), Parts too larga for mounting on a herbarium sheet can be reduced in ize before drying, For instance a large compound palm leaf must be cut up in parts, the number of leaflets counted and cut off on one side of the rhachis (see Drans- field 1986), The fleshy stem of a Hydnophytum can be sliced. Large fruits can beat be dried separately or collected in alcohol, It is ad- visable to cut the fruit lengthwise and across to show the internal structure (number of cells, arrangement of t eds, presence of an aril etc.) and to put the parts in a nylon net, The flowers of some groups of plants are ao delicate that their structure is difficult to reconstruct from herbarium material, e.g. Impatiens, Orchids, Zingi- beraceae, It is best to preserve (part of) the flowers in alcohol 708 or FAA. T£ you collect fallen plant parts try to make sure from which plant they have come, Always mention on your label that the material was picked up from the ground! Certain groups of plants require special collecting techniques. They can not be treated in this brief survey and reference is made to some published instcuctione: arace Nicolson (1965) Balsaminaceae Grey-Wilson (1980) Barbusoidene HcClure (1965), Soderstrom & Young (1983) Lentibulariaceae Taylor (1977) Musaceae Posberg & Sachet (1965) Palmae Dransfield (1974, 1986) Pandanaceae Stone (1983) Pteridophytes Holttum (1957), Henty (1976) Zingiberaceae Burtt & Smith (1976) The special techniques described for collecting members of these groups can also be applied to cther, more or less similar, groups, For instance the instructions for collecting Zingiberaceae also hold for Marantaceae, those for Balsaminaceae are also applicabla to other her- baceous groups such as Begoniaceae, Geuneriaceae and certain Rubiaceae, Tt is recommended always to collect more than one set, The extra ts can be sent to specialists or used as exchange material. Deposit~ 9 your collections in several herbaria is also a kind of inaurance against disasters. Whenever possible, supplement your herbarium collection with 15woodsamples (preferably a part of the main stem) and with pickled material (alcohol 70% or PAA) of flowers and fruits. Always make sure that material of one collecting number comes from one plant, or in case of small plants from one population. If this ie wot possible men- tion it on your label, Make sure that your woodsample and alcohol col- lection gets the same collection number as your herbarium specimen, An opportunity to collect living plants must not be missed. Small plants such as most epiphytic orchids and ferns can simply be wrapped in newspaper and mailed in cartons (cardboard boxes), If it is not possible to send the plants straight away they can be hung on atrings in a shaded place before packing. Sesdlings and terrestrial plants should be collected with ample soil around the rootlets and put in a small plastic bag and tightly packed in newspaper to prevent drying out and damage to the rootlet:. Fruits and seeds can be it in moss, wrapped in newspaper or nylon net. It is advisable to remove all fleshy parte to prevent infection by fungi, Living ferns can be col~ lected by cutting off part of the rootstock, Spores can be collected by placing sori bearing part of the leaf on a newspaper, fold and it. When collecting cuttings, remove the lower leaves, wrap the cuttings in moss and newspaper and put it in a perforated polythene bag with the leaf bearing top sticking out, To promote the development of roots dust the lower end of the cutting with rhizopon, if avail- able. Living material should always be sent back as soon as possible, MAKING NOTES Make it a habit not to rely on your memory but to write your notes at the collecting site ar’ not to wait till the end of the day, or worse still, to put it off to the next day. After collecting your specimen give it a number tag and enter notes in your collection book under the corresponding number. It ig important to take down all par- ticulare that disappear or are invisible in the herbarium specimen (see also van Steenis 1977). Notes should include: a) Location (preferably with reference to a place map, always with latitude and longitude), number. b) Habitat, soil, slope, altitude, vegetation type, etc. c) Habit, tree, shrub, herb, epiphyte, Liana etc. Especially in tree height, d(iameter) (at) b(reast) h(eight), clear bole, shape o! crown, bark characteristics (fiesured, smooth, scaly, colour of living bark, etc.), special root structures, (buttresses, stilt- roots etc.). For description of trees consult Wyatt Smith (1954) and Whitmore (1972), 4a) Exudates latex, resin, (chai of) colour, etc, @) Colour of various parts, such as flowers and fruits and smell of flowers, fruits, crushed leaves etc, £) Ecological and biological objervations: frequency, dominance, pollination of flower: Gispersal of fruits by wind wa! animale, etc, g) Local names, only if reliable (check your informant by showing the same plant a few weeks later and see if he comes up with the same = name). h)- Local us e.g. fruits or leaves edible, bark used for binding, roots used for medicinal purposes etc, Also make notes on Preparation, e.9. seede poisonous raw but edible after cooking, leaves pounded and applied against ulcers, eto, ily located on a late and collection 16Some collectorn prefer to make use of preprinted labels on which the appropriate features are ticked off. This saves a lot of time and makes sure that nothing is forgotten, PRESERVATION IN THE PIELD After making a sample of a plant the procedures for preservation followed deyend on various factors such as time, money, stance available, etc, Thera are two methods of preparing a herbarium specimen: directly drying in the field or provisionally preserving ti material on alcohol anc drying them later in the herbarium (wet mathod) . 4) Drying in the field, ‘The best herbarium speciiens are obtained by drying them as soon as possible after picking. Some prefer to put their specimens directly batween newspaper shi and placa them directly in a plant press (aa- #ag). Others put their specimen in a polythene bag and keep these in a large collection bag (each collection should be placed in a separate bag to avoid mixing of material), Back in camp the material is placed between newspaper sheets, See to it thet the material is well stretch- ed and that part of the 1 are pressed with their underside up- ward. The sheets with corrugates (cardboard or aluminium in between. are made into packs held together with pri and straps. Remember to put number tags with every specimen and/or write the num- ber on the newspaper. Use markers with permanent ink (not dissolving in water or in alcohol). See to it that no parts are sticking out of the bundle and that your material ig evenly distributed. Fill empty spaces with plugs of newspaper. Tha pack is then placed over a source of heat (charcoal or wood fire, kerosene stove, or gas burner) in such away that the hot air is forced to pass through the pack. From time to time check the plant: take out dry specimens and refasten th strape to maintain strong pressure on the specimens (if not the 1 will curl up). The disadvantaga of drying in the field !s that a stove and drier have to be carried or built, It is often difticul< to keep the finished specimens dry and the material takes up a lot of space. b) The "wet" method. A much applied method is to preserve the collected material in alcohol (methylated alcohol or spiritus will also do). This is a time- and space-saving method. The disadvantage is that the packs are heavy and that material treated with the wet method invariably looks dull and unattractive compared to material dried directly. Specimens belonging to one coliection number are wrapped in one or more newspaper sheet eral of these sheets are tied together in bundles and these are placed in a medium sized polythene big. Alcohol ig then poured into the bundle, Use so much alcohol that the newpaper is completely saturated. The bag is then carefully sealed with tape and a number of these bags are stowed into a large polythene bag which in turn is aleo tightly sealed. Make eure that the material is evenly distributed and that no eharp parte stick out and pierce the polythene bags. The alcohol would then evaporate allowing the growth of molds, Material carefully packed in this way can be stored in boxes or crates foc months, It is unpacked and processed later back in the her- barium, 17LITERATURE Burt: BL, & R.M. Smith, 1976, Notes un the collection of Zingibera- ceae, Fl. Hal. Bull, 29: 2599-2601. Dransfield J., 1974. A short guide to rattana, Biotrop 155/74/128: 48-50. -~--, 1986. A guide to collecting palms, Ann. Mo. Bot. Garden 73: 166- 176. Fosberg F.R. & M.H. Sachet, 1965. Manual for tropical herbaria, Regnum Vegetabile 39: 132 pp. Grey-Wilson C., 1980. Notes on co?lecting %mpatiens, Pl, Mal, Bull. 334 3435-3436. Benty &. 1976. Notes on the collection of ferns, in Womersley, l.cet 71. Rolttum R.E., 1957. Instructions for collecting tree ferns ... Pl. Mal. Bull, 13: 567. Hyland B.P.M., 1972. A technique for collecting botanical specimens in rainforest, Pl. Mal, Bull, 261 2038-2040. Jacobs M., 1976. The study of lianas, Pl. Mal. Bull. 29: 2610-2618, McClure P.A,, 1965. Suggestions on how to collect Bamboos, in Fosberg & Bachet l.c. 120-122. Nicolson D.H., 1965. Collecting Avaceae, in Fosberg ¢ Sachet l.c. 123- 126, Perry D.R., 1978, A method of access into the crowns of emergent and canopy trees, Biotropica 10: 155-157. Soderstrom T.R, & 8.M. Young, 1983, A guide to collecting Bamboos, Ann, Mo. Bot. Garden 70: 126-136. Steenis C.G.G.J. van, 1950, The technique of plant collecting and preservation in the tropics, Pl. Mal. I, 11 xlv-lxix. ----, 1977, Three pleas to collectors ..., Fl. Mal. Bull, 30: 2843- 2844, Stone B.C. 1983, Collecting Pandanaceae, Ann, Mo. Bot. Garden 70: 137- 145, Taylor P., 1977, On the collecting and preparation of Utricularia specimens, Fl. Mal, Bull. 30: 2831-2832, Whitmore T.C,, 1972. The description of a tr 29. Womersley J.S., 1976, Plant collecting for Anthropologists, Geog- xaphers and Ecologists in Papua New Guinea, Bot. Bull. no. 2 (2nd ed.)i 75 ppe Womersley J.8., 1981, Plant Collecting and herbarium development, FAO Plant production and protection paper 33. xi + 137 pp. Rome, Wyatt Smith J., 1954, Suggested definitions of field characters, Mal, For, 17: 170-183. Tree Fl, Mal. 1: 23- CHECKLIST OF COLLECTING EQUIPMENT (essential items underlined) @) spotting, measuring and recording binocul binoculars notebook handlens soft black pencils altimeter markers with permanent ink teee-altimeter camera compass taperecorder measuring tape tags/labels 18b) sampling Becateura/clippers catapult + accessories Parang/machete working gloves pocket knife emall aaw pruner c) preserving and sturing Rewspaper polythene. bage/tubing corrugates plastic vials/glass bottles fieldpress plastic containe: adhesive tape nylon wire netting rope + straps alcohol/ FAA canvas bags (methylated) alcohol/spiritus stove small paper bags cartons (cardboard boxes) 19X41, MANUAL FOR THE DESCRIPTION OF FLOWERING PLANTS J.P. Valékamp SUMMARY Practical plant taxonomy is impossible without a thorough know- ledge of and experience in the art of makiing descriptions, The aim of thie manual is to be a guide and companion for beginnere and advanced scientists alike showing the logical sequence of the characters and the terns to be employed, It has been based of manuals used at Dutch Universities for about half a century written by specialists ae Dr. A.A. Pulle, J. Lanjouw, P.A. Plorechetz, and many others to which tho experience of guiding many students during a great number of courses in advanced plant taxonomy at the Rijksherbarium, Leiden, have been INTRODUCTION It is not difficult to describe a plant: you just write down what you see, It is more difficult, at first, to make a good descrip- tion, though. A consequent method must be followed whereby descrip tions are made comparable, while a generally understood vocabulary must be employed to make the intentions clear. Althougn at present English has become a nearly universal scientific language in plant taxonomy it is sucprising to note that there are very few manuals in that language that include practical instructions in phytography, as the subject is called, and all of these are at present difficult to obtain, Apparently students and specialists dc as they please and the results show the lack of a uniform system. It is quite likely that by the absence of a guide to fall back on, many feel insecure and do not like to make descriptions, not knowing where to start, to put what where, and what to call the various forms observed. Yet plant taxonony is based on descriptions: specimens, species, genera, families have to be compared in order to obtain identifications, to study relation shipe, or to obtain a reasonably dependable system, be it based on outward appearances ('phenetic’) or on actual contemporary and his- torical genetic relationships ("phylogenetic') relationships, So when the foundations are weak, how strong can a house be? : Phytography sentially a subjective art. One describes some- thing by its appeararices: if something is large, {iat and green and resembles a leaf, it is called a leaf, Only when it is known through theoretical deductions by specialists in the group involved that it is actually something flattened branch that has taken ov the function of a leaf, may one speak of a phyllodium or phylloclad but it certainly will be confusing for « non-specialist and such definitions only understood by an insider should be avoided. When a description is made a certain sequence and terminology is @asential, In this way the organs have a certain place within the description, whereby they can easily be found back, while the various characters and their variability have a certain place in the sentence. For all parts the dimensions should be given in the appropriote dimen- sions, mm, cm, m. Although the length of a leaf of 110 mm is the same ae 11 om, the former suggests a greater exactness and the latter should be preferred, on the other hand thé length of a sepal of 11 ma da to be preferred over 1.1 cm, which later figure suggests a greater 20measure of fault, When aizes are given one traditionally first give! the length, then the width, then the thickn therefore 6-9 by 3-5 mm is always length by width, even when the firat dimensions are smal- ler than the second: 3-5 by 6-3 mm meane less long than wide. When the dimensions before the 'by' are of the same ‘ 2a as those after it, it is not mentioned: therefore not 6-9 mm b, 3-5 mm, but 6-9 by 3-5 mm, while 6-9 cm by 3-5 mm is correct, Do not use 'x' between such figures, but 'by', it makes a text more easy to read, Colours derived from dry material, as by necessity is usual the case, should be followed by '(i.8.)' (# in sicco, when dried), when taken from field notes or personal observations by '(i.v.)" (= in vivo, alive). In older texte one may find remarks as 'v. (vidi sicco = seen when dried) and 'v.v.' (vidi vivo, geen alive). GOLDEN RULES 1, Be concise. 2, Work from base to top, from outside to inside, from the total to the details. 3. Always try to begin a sentence with its subject. Descriptions should be logical and short. Logical means that one should start to describe the whole and then the parts, that one should begin at the base and finish at the top, that one should work from the outside towards the inside, Short means that all unnecessary words should be avoided, the text should be like a telegram, where each word has to (be paid for. Therefore it is ailly to stater This is a climbing plant. The seeds are red and the leaves are hairy on one side and green on the other, while thore are ten stamens inserted on the blue corolla. There are no stipules, but there are five sepals to the calyx, which have an acute apex and the bark is brown, This shrub is perennial and has leaves which have serrate mar- gins and an acute base and are oblong. Little sense can be made of this, instead it should have been: Climbing shrub. Bark brown, Leaves oblong, base acute, margins serrate, green abova, densely pilose underneath, Stipules absent. Sepals 5, acute, Petals blue. Stamens 10, inserted in the corolla tube. Testa red. There are three types of descriptions: analytical ones (or analyses), diagnostic ones (or diagnoses), and bad ones. dnalytical description: Here everything observed has. been noted. Tt 1s wise to start this way when you are dealing with a group you are unfamiliar with. In this way you will, hopefully, miss nothing essen- tial, Characters belonging to #11 taxonomic levels are included: those pertaining to the family, to the genus, to the species, and to the individual itself, Only after more specimens have been studied of this and other species will you ‘know what is essential and what ia not. It saves more time when you have to cross out what you have writ- 21ten than to go back to thé specimens already studied and to have to add what you have not written dow the first time. Remember: Material is precious! Use as little as possible Diagnostic description: Only those characters by which the taxa @'ffer are noted, Thig can of course only be done when you know the geoup concerned very well, so you should start with an analytical description, Bad description: Characters are haphazardly noted. You, of cour- se, would never make one like that, but many others do. HOW TO START Let's suppose you are starting to revise an entirely new group. You have read up a bit on the family, genus, and the species, but you have never actually studied any material before. First: Carefully (1) inspect the specimens, starting with the species of which there appears to be the most. (If you start with that one you will have probably ‘the most common species, and/or the most variable one, and/cr the one best known to previous collectors and/or the one with the most misidentifications, This last remark brings us to the following: WARNING Names on labels are often wrong! No identification is to be trusted! All humans, including specialists, make mistakes! There is possibly more misidentified than correctly named material! With this in mind sort out the material, Anything vacuely look- ing ont. wf place is to be put aside (Wrong identifications? Aberrant specimens?). Arrange by area, for instance Sumatra, Malaya, Java, @tc., check eagh again for apparent uniformity, check for type material, and then make an analytic description of the most complete and best-looking specimen either of the area where the type material was originally collected, or of the largest pile of material, All being equal, a specimen with inadequate labeling, or coming from a botanical garden should be preferred, A living specimen would of course be best, but that will racely be available. It will often not be possible to obtain all information of all life stages from a ingle specimen, as flowers and truite will be peasent at different times of the year. This is not important, charac ters of any taxonomic value may be derived from apecimens at various stages of the life cycle, The final description will be the sum of the thorough study of all specimens available, Parts of the sum can gradually be collected and added up. ' First of all note the collector's number, (See the pertinent chapter), Alwaya note it on all desoriptions and drawings pertaining to information retrieved from specimens. 22e The description should bo made while ‘following the scheme presented in this manual, It roughly consists of two parts: a vegeta~ ‘tive ’gne and a generative 2 eect one, After the first has, heen nee ‘ * pott a single Flower! _ : ‘you can always boil another one if” required, Fresh material is often easier to handle after boiling, too, Boiling removes the air from dead tissues, or removes the turgor (pressure) within’ living cells. site : aie - "Bring the object in some water in a small beaker, the cap of a whisky or gin bottle is often excellent. Be careful of open firs in an herbarium) Boil it shortly, usually it is enough to have the water boiling for a few seconds, only, after which it may be removed to a Petri-dish ith aufficient water do that it will neither float away, nor be ob- structed by the surface of the water, A drop or two of detergent (in the petridish, not in the beaker) will drive off air bubbles (chaffy “flowers” as in Cyperaceaa, Gramineae do not need to be boiled at all, some detergent in water is eufficient); soak overnight in strong com- mercial ammonia when the floral parts are flimsy and stick together, ae in Balsaminaceae and Orchidaceae, Take care, this is very strong, c. 40%: transfer the object to plain -water,.don't inhale the chemical and see to it that you don't rub it into your eyes with your fingers! Plante cannot count very well, and so, for instance, may produce an occasional 4~, 6-, or even 7-merous flower among the normal 5- merous ones, It will be obvious that somehow one always picks out the odd one out for one's analysis, while descriptions should show the normal condition, $0, before boiling, do check, if possible, that you have not taken an exceptional flower. In case of profusely flowering specimens you can of course afford to boil more than one flower, if you really think you have to. HOWTO PROCEED After the first analytic description has been made two options are open. You may want to finish the description of the taxon, before you atart on the next one, It ia also possible that you are curious to know what the other taxa look like. Some people advocate that you should concentrate on one subject at a time, others say that you will understand your material better if you make a survay first. As the for the actual descriptive method this does not matter much, let's suppose you will want to do the taxa one by one. Aftér the analytic description has been made, take another specimen, preferably originating from a place as close as possible to the first. In this way you may be fairly certain that it is indeed the same species and perhaps may even belong to the same population and there is a chance that variability within a single population may be discovered, Again, first note the collector's number. Now, starting on a new sheet of paper tote only those characters in which this specimen dif- fers from the first. Do not add anything yet to the first original description, why? Suppose you discover that you have two taxa after You will then never be able to reconstruct what information was 23