Introduction

A HANDBOOK OF SYSTEMATIC BOTANY Some Opinions of the Press on " Warming's Systematic Botany." A " very complete, systematic classification of plants, with all their principal characteristics. It is a book well worth reading, and is well printed and illustrated." Lancet. "As a text book of Botany this will take a high place, and as a handbook of reference it will prove valuable. The volume is richly illustrated, and is altogether a serviceable text book for students." Knowledge. " Professor Warming's contribution to morphology and systematic botany are highly valued by botanical workers, who will welcome this translation of his Handbook of We Systematic Botany. commend the book to the notice of those who require to obtain a good general knowledge of the application of morphology to purposes of classification." ttketuettvtt " It has been well translated, and will be found a useful and trustworthy handbook." -Journal of Education. " Mr. Potter has done English students of botany a great service in giving them an excellent translation of Dr. Warming's important work." Educational Review. A HANDBOOK OF SYSTEMATIC BOTANY BY DR. E. WARMING Professor of Botany in the University of Copenhagen WITH A REVISION OF THE FUNGI BY DR. E. KNOBLAUCH, Karlsruhe TRANSLATED AND EDITED BY M. C. POTTER, M.A. F.L.S. Professor of Botany in the University of Durham College of Science, Newcastle-upon- Tyne Author of "An Elementary Text-book of Agricultural Botany WITH 610 ILLUSTRATIONS XonDon SWAN SONNENSCHEIN & CO. LIM NEW YORK : THE MACMILLAN CO 1904 FIRST EDITION, March 1894 ; SECOND EDITION, July 1904 PBEFACE. THE present translation of Dr. E. Warming's Haandbog i den Systematiske Botanik is taken from the text of the 3rci Danish Edition (1892), and from Dr. Knoblauch's German Edition (1890), and the book has been further enriched by numerous additional notes which have been kindly sent to me by the author. Dr. Warming's work has long been recognised as an original and important contribution to Systematic Botanical Literature, and I have only to regret that the pressure of other scientific duties has delayed its presentation to English readers. Dr. Warming desires me to record his high appreciation of the careful transla- tion of Dr. Knoblauch, and his obligation to him for a number of corrections and improvements of which he has made use in the 3rd Danish Edition. In a few instances I have made slight additions to the text ; these, however, appear as footnotes, or are enclosed in square brackets. In the present Edition the Thallophytes have been revised and rearranged from notes supplied to me by Dr. Knoblauch, to whom I am indebted for the Classification of the Fungi, according to the more recent investigations of Brefeld. The Bacteria have been revised by Dr. Migula, the Floridese rearranged after Schmitz, and the Taphrinaceae after Sadebeck. The main body of the text of the Algse and Fungi remains as it was originally written by Dr. Wille and Dr. Rostrup in the Danish Edition, though in many places considerable alterations arid additions have been made. For the sake of comparison a tabular key to the Classification adopted in the Danish Edition is given in the Appendix. In the Angiosperms I have retained the sequence of orders in the Danish original, and have not rearranged them according to w. B. ' * VI PREFACE. the systems more familiar to English students. In any rearrangement much of the significance of Dr. Warming's valuable and original observations would have been lost, and also from a teacher's point of view I have found this system of great value. Although at present it may not be completely satisfactory, yet as an attempt to explain the mutual relationships, development and retrogression of many of the orders, it may be considered to have a distinct advantage over the more artificial systems founded upon Jussieu's Divisions of Polypetalae, Gainopetalse, and Apetalae. With reference to the principles of the systematic arrangement adopted, I may here insert the following brief communication from the author (dated March, 1890), which he has requested me to quote from the preface of Dr. Knoblauch's edition : " Each form which, on comparative morphological considerations, is clearly less simple, or can be shown to have arisen by reduction or through abortion of another type having the same fundamental structure, or in which a further differentiation and division of labour is found, will be regarded as younger, and as far as possible, and so far as other considerations will admit, will be reviewed later than the ' simpler,' more complete, or richer forms. For instance, to serve as an illustration : EPIGYNY and PERIGYNY are less simple than HYPOGNY ; the Epigynous Sympetalse, Chori- petalse, Monoctyledones are, therefore, treated last, the Hydro- charitacecK are considered last under the Helobiece, etc. ZYGOMORPHY is younger than ACTINOMORPHY ; the Scitaminece and Gynandrce therefore follow after the Liliiflorce, the Scrophulariacece after the Solanacece, Linaria after Verbascum, etc. FORMS WITH UNITED LEAVES indicate younger types than those with free leaves ; hence the Sympetalce come after the Choripetalse, the Silenece after the Alsinese, the Malvacese after the Sterculiacese and Tiliacece, etc. "ACYCLIC (spiral-leaved) flowers are older than cyclic (verticillate- leaved) with a definite number, comparing, of course, only those with the same fundamental structure. The Feromca-type must be considered as younger, for example, than Digitalis and Antir- PREFACE. Vii rhinum, these again as younger than Scrophularia ; Verbascum, on the contrary, is the least reduced, and therefore considered as the oldest form. Similarly the one-seeded, nut-fruited Ranunculacece are considered as a later type (with evident abortion) than the many-seeded, folicular forms of the Order ; the Paronychiece and Chenopodiacece as reduced forms of the Alsinece type ; and the occurrence of few seeds in an ovary as generally arising through reduction of the many-seeded forms. The Cyperacece are regarded as a form derived from the Juncacece through reduction, and associated with this, as is so often the case, there is a complication of the inflorescence ; the Dipsacacece are again regarded as a form proceeding from the Valerianacece by a similar reduction, and these in their turn as an off-shoot from the Caprifoliacece, etc. Of course these principles of systematic arrangement could only be applied very generally ; for teaching purposes they have often required modification." In preparing the translation considerable difficulty has been experienced in finding a satisfactory rendering of several terms which have no exact equivalent in English. I may here especially mention the term Vorblatt (Porblad) which I have translated by the term bracteole, when it clearly applied to the first leaf (or leaves) on a pedicel; but in discussing questions of general morphology a term was much needed to include both vegetative and floral shoots, and for this I have employed the term " Foreleaf." Also, the term " Floral-leaf " has been adopted as an equivalent of "Hochblatt," and the term "bract" has been limited to a leaf subtending a flower. At the end of the book will be found a short appendix giving an outline of some of the earlier systems of Classification, with a more complete account of that of Hooker and Bentham. In a book of this character it is almost impossible to avoid some errors, but it is hoped that these will be comparatively few. In correcting the proof-sheets I have received invaluable assist- ance from Dr. Warming and Dr. Knoblauch, who have kindly read through every sheet, and to whom I am greatly indebted for Vlll PREFACE. many criticisms and suggestions. 1 have also to thank Mr. I. H. Burkill for his kind assistance in looking over the proofs of the Monocotyledons and Dicotyledons, and Mr. Harold Wager for kindly reading through the proofs of the Algse and Fungi. My thanks are also especially due to Mr. B. L. Danielsen, and I wish to take this opportunity of acknowledging the very considerable help which I have received from him in translating from the Original Danish. M. C. POTTER. January, 1895. TABLE OF CONTENTS. BEING THE SYSTEM OF CLASSIFICATION ADOPTED IN THE PRESENT VOLUME. (The Algce and Fungi re-arranged in co-operation with Dr. E. Knoblauch, the other Divisions as in the 3rcZ Danish Edition.} ... PAGE DIVISION I. THALLOPHYTA 4 A. Sub-Division. Myxomycetes, Slime-Fungi 5 B. Sub-Division. Algae. Class 1. SYNGENETICJE . . 8 . . . . . .14 ,, 2. DlNOFLAGELLATA 16 ......... 3. DlATOME^E . 18 ,, 4. SCHIZOPHYTA 22 Family 1. Schizophycese 22 2. Bacteria 26 , Class 5. CONJUGATE 41 ,, 6. CHLOKOPHYCEJE 46 ....... Family 1. Protococcoideaj 47 ,, 2. Confervoideee 53 3. Siphoneae 59 .... Class 7. CHAKACE^; 64 ,, 8. PH;EOPHYCE.E (OuvE-BuoAVN SEAWEEDS) 68 Family 1. Phseosporeae 68 ,, 2. Cyclosporero 73 Class 9. DICTYOTALES 76 ,, 10. RHODOPHYCE^E (RED SEAWEEDS) 77 Family 1. Bangioideae ' 2. Floridese . . . " C. Sub-Division. Fungi . Class 1. PHYCOMYCETES . ' Sub-Class 1. Zygomycetes 2. Oomycetes . Family 1. Entomophthorales 2. Chytridiales . 77 ' . . . . .78 .84 96 " 96 ..... 100 102 . . . . .102 3. Mycosiphonales . i . 104 TABLE OF CONTENTS. Class 2. MESOMYCETES ... PAGE 108 Sub- Class 1. Hemiasci 108 2. Hemibasidii ........ Class 3. MYCOMYCETES (HiGHEB FUNGI) Sub- Class 1. Ascomycetes Series 1. Exoasci 109 114 114 116A 2. Carpoasci 118 Family 1. Gymnoascales 118 2. Perisporiales 119 ,, 3. Pyrenomycetes 125 4. Hysteriales 132 5. Discomycetes 132 6, Helvellales . 136 Ascolichenes 136 Sub-Class 2. Basidiomycetes 144 Series 1. Protobasidomycetes 145 2. Autobasidiomycetes 157 Family 1. Dacryomycetes 159 2. Hymenomycetes 159 3. Phalloideje 172 ,, 4. Gasteromycetes 173 Basidiolichenes 176 Fungi Imperfect! 176 DIVISION II. MUSCINE^E (MOSSES) Class 1. HEPATIC^ Family 1. Marchantiese ,, 2. Antboceroteee ,, 3. Jungermanniesa Class 2. Musci FRONDOSI Family 1. ,, 2. 3. ,, 4. Spbagnese Schizocarpese Cleistocarpeae Stegocarpese DIVISION III. PTERIDOPHYTA . Class 1. FILICIN.E Sub- Class 1. Filices Family 1. Eusporangiatee ,, 2. Leptosporangiatsa Sub-Class 2. Hydropteridece Class 2. EQUISEIIN.E (HORSETAILS) . Sub Class 1. Isosporous Equisetince ,, 2. Heterosporous Equisetincc Class 3. LYCOPODINJE (CLUB MOSSES) Sub- Class 1. Lycnpodlea 2. Selagmdlece ... . . 181 188 190 191 191 192 193 195 , t 195 195 . .198 205 205 210 212 215 221 221 . 225 226 22G 228 TABLE OF CONTENTS. XI PAGR TRANSITION FROM THE CRYPTOGAMS TO THE PHANEROGAMS . . . 234 Asexual Generation of the Cormophytes 234 Sexual Generation Fertilisation ; 243 ......... DIVISION IV. GYMNOSPERM^E . . .251 Class 1. CYCADE^E (CYCADS) 252 2. CONIFERS (PINE-TREES) 255 Family 1. Taxoideaj 259 2. Pinoidea 262 Class 3. GNETE^I 270 Fossil Gymnosperms 271 DIVISION V. ANGIOSPERM^E . . .273 Class 1. MONOCOTYLEDONES Family 1. Helobiera 2. Glumiflorffl 3. Spadiciflorse 4. Enantioblast89 5. Liliifloraa 6. Scitaminea 7. Gynandrse Class 2. DICOTYLEDONES Sub-Class 1. Choripetalce Family 1. Saliciflorffi 2. Casuariniflone 3. Querciflorae ,, 4. Juglandiflorsa 5. Urticiflorffi ,,, ,, ^,,, 6. Polygoni floras 7. Curvembryaa 8.' Cactifloree 9. Polycarpicea 10. Ehoeadinse 11. Cistiflor ,, 12. Gruinales ,, 13. Columniferra 14. Tricoccse 15. Terebinthina? 16. Aesculinas 17. Frangulin89 18. ThymelaeineD 19. Saxifraginee 20. EosiflorssK 21. Leguminosse/ 22. Passiflorin 23. MyrtifloraB ^ 24. UmbelliflorsB/ 25. Hysterophyta 274 .278 .283 297 308 309 323 328 334 337 337 339 340 349 351 358 363 375 377 393 406 . .416 421 430 435 439 443 448 451 456 466 475 482 490 498 xu TABLE OF CONTENTS. Sub-Class 2. Sympetalce . A. PentacycliccK . Family 26. Bicornes . 27. DiospyrmsB 28. Primulinas P. Tetracyclicce . Family 29. Tubiflorae 30. Personates 31. Nuculiferse 32. Contort . 33. Kubiales . ,, 34. Dipsacales 35. Campanulinse . 36. Aggregates APPENDIX INDEX PAGE 504 506 506 510 511 514 514, 517 531 541 548 556 560 564 574 593 COEEIGENDA. Page 9, line 12 from top, for Hydrodicton read Hydrodictyon. ,, 14, lines 1 and 2 from top, for as in the- preceding case read in this case. ,, 14, ,, 2 and 15 from top, for zygote read oospore. For a, 6 and ii read ee, 03 and ue throughout. The following are not officinal in the British Pharmacopoeia : page 316, Draccena (Dragon's-blood), Smilax glabra ; p. 321, " Orris-root " ; p. 326, species of Curcuma, Alpinia officinarum ; p. 333, Orchis-species (" Salep "). On page 206, par. 4, only Pearl Barley is offic. in the Brit. Phar. CLASSIFICATION OP THE VEGETABLE KINGDOM. The Vegetable Kingdom is arranged in 5 Divisions. Division I. Thallophyta, Stemless Plants, or those which are composed of a " thallus," i.e. organs of nourishment which are not differentiated into root (in the sense in which this term is used among the higher plants), stem, or leaf. Vascular bundles are wanting. Conjugation and fertilisation in various ways; among most of the Fungi only vegetative multiplication. In contradistinction to the Thallophytes all other plants are called " Stemplants '.' (" Cormophyta "), because their shoots are leaf-bearing stems. The name Thallophyta (Stemless-plants) is to some extent unsuitable, since many of the higher Algae are differentiated into stem and leaf. The Thallophytes are again separated into 3 sub-divisions, namely : Sub-Division A. Myxomycetes, Slime-Fungi, with only 1 class. Sub-Division B. Algae, with 10 classes: Class 1. Syngeneticae. 2. Dinoflagellata, Peridinea. 3. Diatomeae, Diatoms. ,, 4. Schizophyta, Fission Algae. 5. Conjugatae. 6. Chlorophyceae, Green Algae. 7. Characeae, Stone-worts. ,, 8. Phaeophyceae, Brown Algae. 9. Dictyotales. 10. Rhodophyceae, -Red Algse. Sub-Division C. Fungi, with 3 classes: Class 1. Phycomycetes. ,, 2. Mesomycetes. ,., 3. Mycomycetes, Higher Fungi. Division II. Bryophyta or Muscineae, Mosses. These have leaf-bearing shoots, but neither true roots nor vascular w. B B 2 CLASSIFICATION OF THE VEGETABLE KINGDOM. bundles. The lowest Mosses have, however, a thallus. Fertili- sation is accomplished by means of self-motile, spirally coiled spermatozoids, through the agency of water. From the fertilised oosphere a " " fruit-body (capsule) with unicellular organs of reproduction (spores) is produced. The spore on germination gives rise to the vegetative system, which bears the organs of sexual reproduction ; and this system is divided into two stages the protonema, and the leaf-bearing plant produced on it. Alternation of generations : I. The protonema and the entire nutritive system which bears the organs of sexual reproduction. II. The capsule-like sporangium, with spores. 2 Classes : 1. Hepaticse, Liverworts. 2. Musci, Leafy Mosses. Division III Pteridophyta or Vascular Cryptogams, Fern-like Plants having leaf-bearing shoots, true roots, and vascu,lar bundles with tracheides and sieve-tubes. Fertilisation as in the Mosses. From the fertilised oosphere the leaf-bearing shoot arises, which bears on its leaves the reproductive organs, the spores, in capsule-like sporangia. From the germination of the spore a small prothallium is formed, which bears the sexual reproductive organs. Alternation of generations : I. Prothallium with organs of sexual reproduction. II. Leaf-bearing shoot with capsule-like sporangia. 3 Classes: 1. Filicinas, True Ferns. 2. Equisetinae, Horsetails. 3. Lycopodinae, Club-mosses. Division IV. Gymnospermse. The vegetative organs are in the main similar to those in the 3rd Division ; special shoots are modified into flowers for the service of reproduction. From the oosphere, which is fertilised by means of the pollen-tube, the leaf -bearing plant is derived ; this passes the first period of its life as an embryo in the seed, and continues its development when the germination of the seed takes place. The organs correspond- ing to the spores of the two preceding Divisions, are called respec- tively the pollen-grain and embryo-sac. The pollen-grains are multicellular ; i.e. they contain an indistinct prothallium. In the embryo-sac a prothallium, rich in reserve material (endosperm),. CLASSIFICATION OF THE VEGETABLE KINGDOM. 3 with, female organs of reproduction, is developed BEFORE FERTILISATION. The pollen- grains are carried by means of the wind to the ovules; these enclose the embryo-sac, and are situated on the open fruit-leaf (carpel), which has no stigma. Alternation of generations : I. Prothallium = Endosperm in ovule. II. Leaf-bearing plant, with flowers which produce the pollensac and ovule. 3 Classes : 1. Cycadeaa. 2. Conifene. 3. Gnetaceae. Division Y. Angiospermse. The members of this group are very similar to those of Division IV. The ovules are, however, encased in closed fruit-leaves (ovary), which have a special portion (stigma) adapted for the reception and germination of the pollen- grains. The pollen-grains are bicellular, but with only a mem- brane separating the two nuclei ; they are carried to the stigma by animals (chiefly insects), by the wind, or by some other means. Endosperm is not formed till AFTER FERTILISATION. Alternation of generations in the main as in the Gymnosperms, but less distinct ; while the sexual generation, the prothallium, with the organs of fertilisation, is also strongly reduced. 2 Classes: 1 1. Monocotyledones. Embryo with one seed-leaf. 2. Dicotyledones. Embryo with two seed-leaves. For a long time the vegetable kingdom has been divided into CRYPTOGAMS (socalled because their organs of reproduction remained for some time undiscovered), and PHANEROGAMS or Flowering-plants which have evident sexual organs. The first three divisions belong to the Cryptogams, and the third and fourth divisions to the Phanerogams. This arrangement has no systematic value, but is very convenient in many ways. The Cryptogams are also known as Spore -plants, since they multiply byunicellular organs (spores), and the Phanerogams in contradistinction are called Seed-plants (Spermaphyta), since they multiply by seeds, multicellular bodies, the most important part of which is the embryo (a plant in its infancy). Mosses, Ferns, and Gymnosperms are together known as Arche.goniatse, since they possess in common a female organ of distinct structure, the Archegonium. 1 See Angiospermae. 4 THALLOPHYTA. DIVISION I. THALLOPHYTA. The thallus in the simplest forms is unicellular; in the majority, however, it is built up of many cells, which in a few instances are exactly similar ; but generally there is a division of labour, so that certain cells undertake certain functions and are constructed accordingly, while others have different work and corresponding structure. Vessels or similar high anatomical structures are seldom formed, and the markings on the cell-wall are with few exceptions very simple. The Myxomycetes occupy quite an isolated position ; their organs of nourishment are naked masses of protoplasm (plasmodia). As regards the external form, the thallus may be entirely without special prominences (such as branches, members), but when such are present they are all essentially alike in their origin and growth, that is, disregarding the hair- structures A which may be developed. shoot of a Seaweed or of a Lichen, etc., is essentially the same as any other part of the plant ; only among the highest Algae (Characeae, certain Siphonese, Sargassum, and certain Ked Seaweeds) do we find the same differ- ences between the various external organs of the plant body as between stem and leaf, so that they must be distinguished by these names. Roots of the same structure and development as in the Seed- plants are not found, but organs of attachment (rhizoids and haptera) serve partly the biological functions of the root. SYSTEMATIC DIVISION OF THE THALLOPHYTES. To the Thallophytes belong three sub-divisions Slime-Fungi, Algae, and Fungi. Formerly the Thallophytes were divided into Algae, Fungi, and Lichens. But this last group must be placed among the Fungi, since they are really Fungi, which live symbiotically with Algae. The Slime-Fungi must be separated from the true Fungi as a distinct sub-division. The Algce possess a colouring substance, which is generally green, brown, or red, and by means of which, they are able to build up organic compounds from carbonic acid and water. The Bacteria, especially, form an exception to the Algas in this respect ; like the Fungi and Slime-Fungi they have as a rule no such colouring material, but must have organic car- bonaceous food ; these plants form no starch, and need no light MTXOMYCETES. 5 for their vegetation (most Fungi require light for fructification). The Myxomycetes, Bacteria, and Fungi derive their nourishment either as saprophytes from dead animal or vegetable matter, or as parasites from living animals or plants (hosts), in which they very often cause disease. A remark, however, must be made with regard to this division. Among the higher plants so much stress is not laid upon the biological relations as to divide them into " green " and " non-green"; Cuscuta (Dodder), a parasite, is placed among the Convolvulaceae, Neottia and Corallorhiza, saprophytes, belong to the Orchidaceas, although they live like Fungi, yet their relations live as Algae. In the same manner there are some colourless parasitic or saprophytic forms among the Algae, and stress must be laid upon the fact that not only the Blue-green Algae, but also the Bacteria, which cannot assimilate carbonic-acid, belong to the Algae group, Sehizophyceae. The reason for this is that systematic classifications must be based upon the relationship of form, development, and reproduction, and from this point of view we must regard the Bacteria as being the nearer relatives of the Blue-green Algae. All the Thallophytes, which are designated Fungi (when the entire group of Slime-Fungi is left out), form in some measure a connected series of development which only in the lower forms (Phycomycetes) is related to the Alga?, and probably through them has taken its origin from the Alg83 ; the higher Fungi have then developed independently from this beginning. The distinction of colour referred to is therefore not the only one which separates the Alga? from the Fungi, but it is almost the only characteristic mark by which we can at once distinguish the two great sub-divisions of the Thallophytes. The first fo>ms of life on earth were probably " Protistae," which had assimilating colour material, or in other words, they were Algae because they could assimilate purely inorganic food substances, and there are some among these which belong to the simplest forms of all plants. Fungi and Slime-Fungi must have appeared later, because they are dependent on other plants which assimi- late carbon. 1 Sri-Division I. MYXOMYCETES, SLIME-FUNGI. The Slime-Fungi occupy quite an isolated position in the Vegetable Kingdom, and are perhaps the most nearly related to the group of Rhizopods in the Animal Kingdom. They live in and on organic remains, especially rotten wood or leaves, etc., on the surface of which their sporangia may be found. They are organisms without chlorophyll, and in their vegetative condition are masses of protoplasm without cell- wall (plasmodia) . They multiply by means of spores, which in the true 2 Slime-Fungi 1 According to the recent investigations of Winogradsky some micro-organisms (Nitrifying-bacteria) can build organic from inorganic matter. Sachs' hypothesis that the first organisms must necessarily have contained chlorophyll is there- fore untenable. 2 Myxogasteres, Engler's Syllabus, p. 1. 6 MYXOMYCETES. are produced in sporangia, but in some others l free. The spores are round cells (Fig. 1 a) which in all the true Slime-Fungi are surrounded by a cell- wall. The wall bursts on germination, arid the contents float out in the water which is necessary for germination. They move about with swimming and hopping motions like swarmspores (e, /), having a cilia at the front end and provided with a cell-nucleus and a pulsating vacuole. Later on FiG.l. a-l Development of "Fuligo" from spore to Myxamceba ; a-m are magnified m 390 times ; is a Myxamoeba. of Lycogala epidendron ; I' three Myxamosbse of Physarum album about to unite; o, a small portion of plasmodium, magnified 90 times. they become a little less active, and creep about more slowly, while they continue to alter their form, shooting out arms in various places and drawing them in again (g, h, i, k, I, tn) ; in this stage they are called Myzamoebce. The Myxamceba grows whilst taking up nourishment from the material in which it lives, and multiplies by division. At a later stage a larger or smaller number of A1yxamceba3 may be seen to 1 Acrasieae and Plasmodiophorales, ibid. MYXOMYCETES. coalesce and form large masses of protoplasm, plasmodia, which in the " Flowers of Tan " may attain the size of the palm of a hand, or even larger, but in most