Part 1

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

London SWAN SONNENSCHEIN & CO NEW YORK: MACMILLAN & CO 1895

BUTLER & TANNER, THE SELWOOD PRINTING WORKS, FROME, AND LONDON.

PREFACE.

The present translation of Dr. E. Warming’s _Haandbog i den Systematiske Botanik_ is taken from the text of the 3rd 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 translation 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 Florideæ rearranged after Schmitz, and the Taphrinaceæ after Sadebeck. The main body of the text of the Algæ and Fungi remains as it was originally written by Dr. Wille and Dr. Rostrup in the Danish Edition, though in many places considerable alterations and 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 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 Polypetalæ, Gamopetalæ, and Apetalæ.

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 _Sympetalæ_, _Choripetalæ_, _Monocotyledones_ are, therefore, treated last, the _Hydrocharitaceæ_ are considered last under the _Helobieæ_, etc. ZYGOMORPHY is younger than ACTINOMORPHY; the _Scitamineæ_ and _Gynandræ_ therefore follow after the _Liliifloræ_, the _Scrophulariaceæ_ after the _Solanaceæ_, _Linaria_ after _Verbascum_, etc. FORMS WITH UNITED LEAVES indicate younger types than those with free leaves; hence the _Sympetalæ_ come after the _Choripetalæ_, the _Sileneæ_ after the _Alsineæ_, the _Malcaceæ_ after the _Sterculiaceæ_ and _Tiliaceæ_, 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 _Veronica_-type must be considered as younger, for example, than _Digitalis_ and _Antirrhinum_, 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 _Ranunculaceæ_ are considered as a later type (with evident abortion) than the many-seeded, folicular forms of the Order; the _Paronychieæ_ and _Chenopodiaceæ_ as reduced forms of the _Alsineæ_ type; and the occurrence of few seeds in an ovary as generally arising through reduction of the many-seeded forms. The _Cyperaceæ_ are regarded as a form derived from the _Juncaceæ_ through reduction, and associated with this, as is so often the case, there is a complication of the inflorescence; the _Dipsacaceæ_ are again regarded as a form proceeding from the _Valerianaceæ_ by a similar reduction, and these in their turn as an offshoot from the _Caprifoliaceæ_, 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 (Forblad) 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 “Fore-leaf.” 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. I have followed Dr. E. L. Mark in translating the word “Anlage” by “Fundament.”

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 assistance from Dr. Warming and Dr. Knoblauch, who have kindly read through every sheet, and to whom I am greatly indebted for many criticisms and suggestions. I 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 Algæ and Fungi. My thanks are also especially due to Mr. E. 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 Algæ and Fungi rearranged in co-operation with Dr. E. Knoblauch, the other Divisions as in the 3rd Danish Edition._)

PAGE

DIVISION I. THALLOPHYTA 4

A. Sub-Division. Myxomycetes, Slime-Fungi 5

B. Sub-Division. Algæ 8

Class 1. SYNGENETICÆ 14

„ 2. DINOFLAGELLATA 16

„ 3. DIATOMEÆ 18

„ 4. SCHIZOPHYTA 22

Family 1. Schizophyceæ 22

„ 2. Bacteria 26

Class 5. CONJUGATÆ 41

„ 6. CHLOROPHYCEÆ 46

Family 1. Protococcoideæ 47

„ 2. Confervoideæ 53

„ 3. Siphoneæ 59

Class 7. CHARACEÆ 64

„ 8. PHÆOPHYCEÆ (OLIVE-BROWN SEAWEEDS) 68

Family 1. Phæosporeæ 68

„ 2. Cyclosporeæ 73

Class 9. DICTYOTALES 76

„ 10. RHODOPHYCEÆ (RED SEAWEEDS) 77

Family 1. Bangioideæ 77

„ 2. Florideæ 78

C. Sub-Division. Fungi 84

Class 1. PHYCOMYCETES 96

Sub-Class 1. _Zygomycetes_ 96

„ 2. _Oomycetes_ 100

Family 1. Entomophthorales 102

„ 2. Chytridiales 102

„ 3. Mycosiphonales 104

Class 2. MESOMYCETES 108

Sub-Class 1. _Hemiasci_ 108

„ 2. _Hemibasidii_ 109

Class 3. MYCOMYCETES (HIGHER FUNGI) 114

Sub-Class 1. _Ascomycetes_ 114

Series 1. Exoasci 116

„ 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. Phalloideæ 172

„ 4. Gasteromycetes 173

Basidiolichenes 176

Fungi Imperfecti 176

DIVISION II. MUSCINEÆ (MOSSES) 181

Class 1. HEPATICÆ 188

Family 1. Marchantieæ 190

„ 2. Anthoceroteæ 191

„ 3. Jungermannieæ 191

Class 2. MUSCI FRONDOSI 192

Family 1. Sphagneæ 193

„ 2. Schizocarpeæ 195

„ 3. Cleistocarpeæ 195

„ 4. Stegocarpeæ 195

DIVISION III. PTERIDOPHYTA 198

Class 1. FILICINÆ 205

Sub-Class 1. _Filices_ 205

Family 1. Eusporangiatæ 210

„ 2. Leptosporangiatæ 212

Sub-Class 2. _Hydropterideæ_ 215

Class 2. EQUISETINÆ (HORSETAILS) 221

Sub-Class 1. _Isosporous Equisetinæ_ 221

„ 2. _Heterosporous Equisetinæ_ 225

Class 3. LYCOPODINÆ (CLUB MOSSES) 226

Sub-Class 1. _Lycopodieæ_ 226

„ 2. _Selaginelleæ_ 228

TRANSITION FROM THE CRYPTOGAMS TO THE PHANEROGAMS 234

Asexual Generation of the Cormophytes 234

Sexual Generation; Fertilisation 243

DIVISION IV. GYMNOSPERMÆ 251

Class 1. CYCADEÆ (CYCADS) 252

„ 2. CONIFERÆ (PINE-TREES) 255

Family 1. Taxoideæ 259

„ 2. Pinoideæ 262

Class 3. GNETEÆ 270

Fossil Gymnosperms 271

DIVISION V. ANGIOSPERMÆ 273

Class 1. MONOCOTYLEDONES 274

Family 1. Helobieæ 278

„ 2. Glumifloræ 283

„ 3. Spadicifloræ 297

„ 4. Enantioblastæ 308

„ 5. Liliifloræ 309

„ 6. Scitamineæ 323

„ 7. Gynandræ 328

Class 2. DICOTYLEDONES 334

Sub-Class 1. _Choripetalæ_ 337

Family 1. Salicifloræ 337

„ 2. Casuarinifloræ 339

„ 3. Quercifloræ 340

„ 4. Juglandifloræ 349

„ 5. Urticifloræ 351

„ 6. Polygonifloræ 358

„ 7. Curvembryæ 363

„ 8. Cactifloræ 375

„ 9. Polycarpicæ 377

„ 10. Rhœadinæ 393

„ 11. Cistifloræ 406

„ 12. Gruinales 416

„ 13. Columniferæ 421

„ 14. Tricoccæ 430

„ 15. Terebinthinæ 435

„ 16. Aesculinæ 439

„ 17. Frangulinæ 443

„ 18. Thymelæinæ 448

„ 19. Saxifraginæ 451

„ 20. Rosifloræ 456

„ 21. Leguminosæ 466

„ 22. Passiflorinæ 475

„ 23. Myrtifloræ 482

„ 24. Umbellifloræ 490

„ 25. Hysterophyta 498

Sub-Class 2. _Sympetalæ_ 504

_A. Pentacyclicæ_ 506

Family 26. Bicornes 506

„ 27. Diospyrinæ 510

„ 28. Primulinæ 511

_B. Tetracyclicæ_ 514

Family 29. Tubifloræ 514

„ 30. Personatæ 517

„ 31. Nuculiferæ 531

„ 32. Contortæ 541

„ 33. Rubiales 548

„ 34. Dipsacales 556

„ 35. Campanulinæ 560

„ 36. Aggregatæ 564

APPENDIX 574

INDEX 593

CORRIGENDA.

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_. „ 88, line 15 from bottom, for _Periphyses_ read _periphyses_. „ 124, „ 7 „ „ for _Chæromyces_ read _Choiromyces_. „ 142, „ 2 „ „ and in Fig. 137, for _Bœomyces_ read _Bæomyces_. „ 152, „ 2 „ top, for _Pirus_ read _Pyrus_. „ 152, „ 5 „ „ for _Crategus_ read _Cratægus_. „ 216, Fig. 215, for _Salvina_ read _Salvinia_. „ 306, line 6 from top, for _Pista_ read _Pistia_. „ 316, „ 26 „ „ after Dracæna insert a comma. „ 337, „ 13 „ „ for _end_ read _beginning_. „ 483, „ 11 „ bottom, for _Lagerstrœmia_ read _Lagerstrœmeria_.

For ä, ö and ü read æ, œ and ue throughout.

The following are not officinal in the British Pharmacopœia:--page 316, _Dracæna_ (Dragon’s-blood), _Smilax glabra_; p. 321, “Orris-root”; p. 326, species of _Curcuma_, _Alpinia officinarum_; p. 333, _Orchis_-species (“Salep”). On page 296, par. 4, only Pearl Barley is offic. in the Brit. Phar.

CLASSIFICATION OF 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 “Stem-plants” (“Cormophyta”), because their shoots are leaf-bearing stems. The name Thallophyta (Stemless-plants) is to some extent unsuitable, since many of the higher Algæ 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.=--=Algæ=, with 10 classes: Class 1. Syngeneticæ. „ 2. Dinoflagellata, Peridinea. „ 3. Diatomeæ, Diatoms. „ 4. Schizophyta, Fission Algæ. „ 5. Conjugatæ. „ 6. Chlorophyceæ, Green Algæ. „ 7. Characeæ, Stone-worts. „ 8. Phæophyceæ, Brown Algæ. „ 9. Dictyotales. „ 10. Rhodophyceæ, Red Algæ.

Sub-Division =C.=--=Fungi=, with 3 classes: Class 1. Phycomycetes. „ 2. Mesomycetes. „ 3. Mycomycetes, Higher Fungi.

Division II.--=Bryophyta or Muscineæ, Mosses.= These have leaf-bearing shoots, but neither true roots nor vascular bundles. The lowest Mosses have, however, a thallus. Fertilisation 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. Hepaticæ, Liverworts. 2. Musci, Leafy Mosses.

Division III.--=Pteridophyta or Vascular Cryptogams=, =Fern-like Plants= having leaf-bearing shoots, true roots, and vascular 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. Filicinæ, True Ferns. 2. Equisetinæ, Horsetails. 3. Lycopodinæ, Club-mosses.

Division IV.--=Gymnospermæ.= 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 corresponding to the spores of the two preceding Divisions, are called respectively 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), 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 pollen-sac and ovule. 3 Classes: 1. Cycadeæ. 2. Coniferæ. 3. Gnetaceæ.

Division V.--=Angiospermæ=. 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 membrane 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 (so called 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 by unicellular 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 Archegoniatæ, since they possess in common a female organ of distinct structure, the Archegonium.

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 which may be developed. A shoot of a Seaweed or of a Lichen, etc., is essentially the same as any other part of the plant; only among the highest Algæ (Characeæ, certain Siphoneæ, _Sargassum_, and certain Red Seaweeds) do we find the same differences 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.