O? m r-R CD m en GRAY'S BOTANICAL TEXT-BOOK. VOLUME I. STRUCTURAL BOTANY. GRAY'S BOTANICAL TEXT-BOOK CONSISTS OF VOL. I. STRUCTURAL BOTANY. By ASA GRAY. II. PHYSIOLOGICAL BOTANY. By GEORGE L. GOODALE. III. INTRODUCTION TO CRYPTOGAMIC BOTANY, BOTH STRUCTURAL AND SYSTEMATIC. By WILLIAM G. FARLOW. (In preparation.) IV. SKETCH OF THE NATURAL ORDERS OF PH^ENOGAMOUS PLANTS ; their Special Morphology, Useful Pro- ducts, &c. (In preparation.) GRAY'S BOTANICAL TEXT-BOOK. (SIXTH EDITION.) VOL. I. STRUCTURAL BOTANY, OK ORGANOGRAPHY ON THE BASIS OF MORPHOLOGY. TO WHICH IS ADDED THE PRINCIPLES OF TAXONOMY AND PHYTOGRAPHY, AND | of botanical Ccrm0. BY ASA GRAY, LL.D., ETC., FISHER PROFESSOR OF NATURAL HISTORY (BOTANY) IN HARVARD UNIVERSITY. NEW YORK •:• CINCINNATI •:• CHICAGO AMERICAN BOOK COMPANY GRAY'S BOTANICAL SERIES Gray's How Plants Grow Gray's How Plants Behave Gray's Lessons in Botany Gray's Field, Forest, and Garden Botany (Flora only) Gray's School and Field Book of Botany (Lessons and Flora) Gray's Manual of Botany. (Flora only) Gray's Lessons and Manual of Botany Gray's Botanical Text-Book 1. Gray's Structural Botany II. Goodale's Physiological Botany Coulter's Manual of Botany of the Rocky Mountains Gray and Coulter's Text-Book of Western Botany Copyright, 1879, by ASA GRAY GRAY'S STRUCTURAL BOTANY w. P. 4 PREFACE. THE first edition of this treatise was published in the year 1842, the fifth in 1857. Each edition has been in good part rewritten, — the present one entirely so, — and the compass of the work is now extended. More elementary works than this, such as the author's First Lessons in Botany (which contains all that is necessary to the prac- tical study of systematic Phsenogamous Botany by means of Manuals and local Floras), are best adapted to the needs of the young beginner, and of those who do not intend to study Botany comprehensively and thoroughly. The present treatise is intended to serve as a text-book for the higher and completer instruction. To secure the requisite fulness of treatment of the whole range of sub- jects, it has been decided to divide the work into distinct volumes, each a treatise by itself, which may be indepen- dently used, while the whole will compose a comprehensive botanical course. This volume, on the Structural and Morphological Botany of Phsenogamous Plants, properly comes first. It should thoroughly equip a botanist for the scientific prosecution of Systematic Botany, and furnish needful preparation to those who proceed to the study of Vegetable Physiology and Anatomy, and to the wide and varied department of Cryptogamic Botany. iv I'KKKACE. The preparation of the volume upon Physiological Botany (Vegetable Histology and Physiology) is assigned to the author's colleague, Professor GoODALE. The Introduction to Cryptogamous Botany, both structu- ral and systematic, is assigned to his colleague, Professor FARLOW. A fou iih volume, a sketch of the Natural Orders of Phsenogamous Plants, and of their special Morphology, Classification, Distribution, Products, &c., will be needed to complete the series : this the present author may rather hope than expect himself to draw up. ASA GRAY. \ HERBARIUM OF HARVARD UNIVERSITY, CAMBRIDGE April 10, 1879. *#* The numerals in parentheses, which are here and there introduced into sentences or appended to them, are references to the numbered para- graphs in which the topic is treated or the term explained. CONTENTS. PAGE INTRODUCTION. THE DEPARTMENTS OF THE SCIENCE .... 1 CHAPTER I. OUTLINES OF THE GENERAL MORPHOLOGY OF PH^ENOGAMOUS PLANTS 5 CHAPTER II. MORPHOLOGY AND DEVELOPMENT OF THE EMBRYO AND SEEDLING 9 The Embryo, its Nature, Structure, and Parts 9 Development of the Dicotyledonous Embryo in Maple ... 10 In Ipomoea, or Morning Glory, &c., with Albuminous Seeds . 13 In Embryos with thickened Cotyledons 16 As of Almond, Beech, Bean, &c 17 With Hypogaeous Germination and no Elongation of Caulicle 19 In Megarrhiza, &c., with concreted Petioles to the Cotyledons 21 In Ipomoea leptophylla with foliaceous and long-petioled Cotyledons and no elongation of Caulicle 22 In Pumpkin, &c., with no Primary Root 22 The Polycotyledonous Embryo 23 The Monocotyledonous Embryo of Iris, Onion, Cereal Grains 24 Pseudo-monocotyledonous and Acotyledonous Embryo ... 26 Dicotyledonous and Monocotyledonous Plants 27 CHAPTER III. MORPHOLOGY AND STRUCTURE OF THE ORGANS OF THE PLANT IN VEGETATION . . 27 SECTION I. OF THE ROOT 27 Nature, Growth, and Composition 28 Root-hairs 29 Kinds of Roots 29 Duration ; Annuals 30 Biennials 31 Perennials 32 Aerial Roots 33 Epiphytes or Air-plants 35 Parasitic Plants, Green and Colored 36 VI CONTENTS. SECTION II. OF Btn>s 40 Scaly Buds and Bud-scales 40 Naked, Snbpetiolar, and Fleshy Buds 41 Bud-propagation 43 Normal, Accessory, and Adventitious Buds 44 SECTION III OF THE STEM 45 § 1. GENERAL CHARACTERISTICS AND GROWTH 45 Development and Structure 46 Ramification, Branches 47 Kxrurrent and Deliquescent Stems 48 Definite and Indefinite Annual Growth 49 §2. FORMS OF STI.M \M> BK \MIIES 50 I lerbs. Shrubs, Trees, Culm, Caudex, Scape 50 Climbing Stems, Twining or otherwise 51 Leaf -( Timbers, Tendril-climbers, and Knot-climbers .... •">:.' Suckers, Stolons, Offsets, Runner- •'•'• Tendrils formed of Stems -M Sympodial and Monopodial Stems 55 Spines or Thorns and Subterranean Stems 56 l.'iiixoma or Rootstock 57 Tuber, Tubercles 59 Conn or Solid Bulb 61 Hull), Bull.le.ts ''•- Condensed Aerial Stems 64 Stems serving for Foliage, Phyllocladia, Cladophylla ... 65 Frondose Stems 66 § 3. INTERNAL STRUCTURE 67 Anatomical KleinenN 68 Endogenous Structure 70 K \ogenous Structure; its Beginning ~:> First Year's Growth "' Pith, Layer of Wood. &c 75 Bark, its Part- and Structure 76 Annual Increase in Diameter 78 Demarcation of Annual Layers 79 Sap-wood and Heart-wood 80 :; Forms as to Outline .... 94 Forms as to Extremity 96 Forms as to Margin or Special Outline and Dentation ... 97 Lobation or Segmentation 98 Number and Arrangement of Parts 99 Compound Leaves, Pinnate and Palmate or Digitate, &c. . . 100 Petiole or Leafstalk t . . . 104 Stipules, Ligule, StipeLs 1<>"> Leaves in unusual Modifications 10(5 Such as Inequilateral, Connate, Perfoliate 107 Vertical and Equitant 108 Without distinction of Parts 109 Stipules serving for Blade 109 Phyllodia, or Petioles serving for Blade llf § 3. LEAVES SERVING SPECIAL OFFICES . Hr Utilizing Animal Matter . 110 Ascidia or Pitchers Sensitive Fly-traps Leaves for Storage H'r> Bulb-scales and Bud-scales CHAPTER IV. PHYLLOTAXY, OR LEAF-ARRANGEMENT . 119 SECTION I. DISTRIBUTION OF LEAVES ON THE STEM .... 119 Phyllotaxy either Verticillate or Alternate, Cyclical or Spiral 119 Verticillate or Cyclical Arrangement 120 Alternate or Spiral Arrangement 121 Its Modes and Laws 122 Relation of Whorls to Spirals 129 Hypothesis of the Origin of Both 130 Fascicled Leaves 131 SECTION II. DISPOSITION OF LEAVES IN THE BUD 132 Vernation and ^Estivation ; the Modes 132 Direction, Dextrorse and Sinistrorse 140 VI 11 CON TK NTS. CHAPTER V. ANTHOTAXV <>1! INFLORESCENCE .... 141 Bracts and Brartlets and their Modifications 141 Peduncles, Pedicels, Rliachis, Receptacle 14-'! Position of Flower-buds, Kinds of Inflorescence 144 Indeterminate, Indefinite, or Botryose 140 Kaeenie. C'oryinh, I'mliel 140 Head or < 'apitulum 147 Syconium or Hypanthodium 148 Spike, Spadix, Ament or Catkin 14'.* Panicle and other Compound Forms 150 Determinate or Cymose . 151 Cyme, Glomerule, «jic. ... ... . . . . Botryoidal Forms of Cymose Type Sympodial Forms 154 Scorpioid and Helicoid, the Pleiochasium, Dichasium, and Monochasium 155 Bostryx, Cincinnus, Rhipidiuiu, Drepanium, «!cc !•")<'> Mixed Inflorescence 158 Thyrsus, Verticillaster, &c 1-V.i Relations of Bract, Bractlet, and Flower 160 Anterior and Posterior, or Inferior and Superior 100 Median and Transverse 1(50 Position of Bractlets 161 Tabular View of Inflorescence 102 CHAPTER VI. THE FLOWER 163 SECTION I. ITS NATURE, PARTS, AND METASIORPHY .... 163 Floral Envelopes, Perianth, or Perigone 164 Tlie Parts, Calyx and Corolla . . 1 ('.."• Aiulra'cium, Stamens 165 Gynoecium, Pistils 160 Torus or Receptacle of the Flower 107 Metamorphosis 167 Unity of Type illustrated by Position and Transitions . . . 169 Teratological Transitions and Changes 170 SECTION II. FLORAL SYMMETRY 174 Symmetrical, Regular, and Complete Flower 175 Niimerieal Ground-plan 17<> Pattern Flowers 170 Diplostemonous Type . 177 SECTION III. VARIOUS MODIFICATIONS OF THE FLOWER . . . 179 § 1. EM MI. i: \ i io\ or mi: KINDS 179 §2. REGULAR UNION OF SIMILAR PARTS 180 Coalescence or Cohesion . 180 CONTENTS. ix § 3. UNION OF DISSIMILAR OR SUCCESSIVE PARTS .... 181 Adnation or Connation 182 Hypogynous, Perigynous, Epigynous 183 § 4 IRREGULARITY OF SIMILAR PARTS 184 § 5. DISAPPEARANCE OR OBLITERATION OF PARTS .... 187 Abortion or Suppression of Parts of a Circle 187 Abortion or Suppression of whole Circles 190! Terms therewith connected 191 Suppressed Perianth 191 Suppressed Androecium or Gynoeciuui 193 Along with suppressed Perianth 194 Neutral Flowers 195 § 6. INTERRUPTION OF NORMAL ALTERNATION 195 Anteposition or Superposition . . .... .... 195 In Appearance only 196 Superposition by Spirals ... 196 Anteposition with Isostemony and Diplostemony 197 With Obdiplosteraony 198 § 7. INCREASED NUMBER OF PARTS 200 Regular Multiplication 200 Parapetalous Multiplication 201 Chorisis or Deduplication 202 § 8. OUTGROWTHS 209 Their relation to Chorisis : Trichomes 209 Corona or Crown 210 Ligule 211 § 9. FORMS OF THE TORUS OR RECEPTACLE 211, Stipe, Thecaphore, Gynophore, Carpophore, &c 212 Disk 213 Hypanthium 214 SECTION IV. ADAPTATIONS OF THE FLOWER TO THE ACT OF FERTILIZATION 215 § 1. IN GENERAL ' . . . . 215 Close and Cross Fertilization, or Autogamy and Allogamy . 216 § 2. ADAPTATIONS FOR ALLOGAMY OR INTERCROSSING . . . 216 Wind-fertilizable or Anemophilous Flowers 217 Insect-fertilizable or Entomophilous Flowers 218 Irregularity as related to Allogamy 219 CONTENTS. Dichogamy, cither Proterandrous or Proterogynous .... 219 Froiei-ou-yny Proterandry Particular Ailaptaiion- in Papilionaceous Flowers .... In Kalmia-blossoms, Iri>. &c Transportation of 1'ullinia In Orchidacese and Asclepiadaceae 231 Heterogonous Dimorphism and Trimorphism L'-'ll §3. ADAPTATIONS i <>u CLOSE FERTILIZATION 240 Clei.-to.i:ainy 241 SECTION V. THE PKIMANTII. OR THE CALYX AND COROLLA IN J'AKTK 1 |. Vlt Perianth as to Duration, Numerical Terms, Union, &c. , . . Parts of Petals and of Gamophyllous Perianth 245 Forms of Corolla and Calyx 246 SECTION VI. Tin: AN-HUCEC-ITM, OR STAMENS IN PARTICULAR . 2H» The Stamen as a whole ; Numerical Terms iM'.i The Filament and the Anther; their Modifications .... 251 Pollen 256 Pollen-tubes 258 SECTION VII. THE PISTILS, OR GTNOECIUM 259 § 1. IN ANGIOSPERMS 259 Carpel or Carpophyll . 260 Ventral and Dorsal Sutures ; 1'lacenta 261 Simple or Apocarpous Pistils -''''-' Compound or Syncarpous Pistil if''-1- With two or more Cells and Axile Placentae ; Partitions . . i-'i'-l With one Cell and Parietal Placentae 265 With one Cell and I-W Central Placenta . '-'<'><'> Anomalous Placentation -(i~ § 2. IN GYMNOSIM.UMS 268 Structure in (Inetacea- -';;l Structure in Coniferse 270 In the Yew Family 271 In the Pine Tribe, &c _ -~- In the C\ pros Tribe 273 Structure in Cycadaiva- 274 SECTION \'I1I. Tin: Ovri.i; 276 Its Structure and Position 277 \\- Forms, ( >rthotropous, Campylotropous, Ampliitropous, Anairojious 278 Origin and Morphological Nature of the Ovule 282 of the Fmbryo 283 CONTENTS. XI CHAPTER VII. THE FRUIT 285 SECTION I. ITS STRUCTURE, TRANSFORMATIONS, AND DEHIS- CENCE 285 Pericarp, its Alterations, Accessions, and Transformations . 287 Dehiscence 288 SECTION II. THE KINDS OF FRUIT 291 Simple Fruits 291 Dehiscent Fruits, Follicle, Legume, Capsule, Pyxis, Silique . 292 Indehiscent Dry Fruits, Samara, Akene, Utricle, Caryopsis, Nut, &c 294 Fleshy Fruits, Drupe, Pome, Pepo, Berry, &c 297 Aggregate Fruits 299 Accessory or Anthocarpous Fruits 300 Multiple or Collective Fruits, Syconium, Strobile, &c. . . . 301 Table of Simple Fruits 304 CHAPTER VIII. THE SEED 305 Its Stalk, Coats, and Appendages 306 Aril or Arillus 308 Nucleus or Kernel, Albumen 309 The Embryo, its Parts and Positions 311 The Cotyledons as to Adjustment and Number 313 CHAPTER IX. TAXONOMY 315 SECTION I. THE PRINCIPLES OF CLASSIFICATION IN NATURAL HISTORY 315 Individuals 315 Species 317 Varieties, Races, &c 318 Cross-breeds and Hybrids 321 Genera 323 Orders, Classes, Tribes, &c 325 Sequence of the Grades 327 Nature and Meaning of Affinity 327 Theory of Descent and Natural Selection 328 SECTION II. BOTANICAL CLASSIFICATION 331 Ante-Linnaean Classifications 332 Linnaean Classification 333 Sexual Artificial System 334 Natural System 338 As presented by Jussieu 339 Some of its Modifications 340 xii CONTENTS. CHAPTER X. PIIYTOGRAPHY 345 SKCTION I. NOMENCLATURE 345 Names of Plants, Binomial Nomenclature 346 Rules for naming Plants -'-IT Names of (ienera ... 348 Names of Species, Varieties, &c ^-'i11 The Fixation, Precision, and Citation of Names Subgeiu ric Names '•'••><< Tribal ami < >nlinal Names 357 Names of Cohorts, Classes, &c 358 SECTION II. GLOSSOLOGY OR TERMINOLOGY 350 SECTION III. DESCRIPTION 801 Characters :>(i' Punctuation ;>*>1 Synonomy 365 Iconography 366 Habitat and Station, &c 366 Etymology of Names 366 Accentuation, Abbreviations 367 Signs 368 Floras, Monographs, &c. ... 369 SECTION IV. SPECIMENS, DIRECTIONS FOE THEIR EXAMINA- TION, PRESERVATION, &c 370 Implements of Investigation 370 Diagrams :>il Herborizing :»71 Drying Specimens •'"•> Poisoning Specimens 379 The Herbarium 380 ABBREVIATIONS 385 SIGNS 391 GLOSSARY OF BOTANICAL TERMS, WITH INDEX . . 393 STRUCTURAL BOTANY ON THE BASIS OF MORPHOLOGY. INTRODUCTION. 1. THE two Biological Sciences,1 considered as parts of Natural History, are Zoolog}* and Botany. The latter is the natural history of the Vegetable Kingdom. It embraces every scientific inquiry that can be made respecting plants, their nature, their kinds, the laws which govern them, and the part they play in the general economy of the world. 2. We cannot distinguish the vegetable from the animal king- dom by any complete and precise definition. Although ordinary observation of their usual representatives may discern little that is common to the two, yet there are many simple forms of life which hardly rise high enough in the scale of being to rank dis- tinctively either as plant or animal ; there are undoubted plants possessing faculties which are generally deemed characteristic of animals ; and some plants of the highest grade share in these endowments. But in general there is a marked contrast between animal and vegetable life, and in the part which animals and plants respectively play in nature. 3. Plants only are nourished upon mineral matter, upon earth and air. It is their peculiar office to appropriate mineral mate- rials and to organize them into a structure in which life is mani- fested, — into a structure which is therefore called organic. So the material fitted for such structure, and of which the bodies 1 Biology, the science of life, or rather of living things, in its earlier use was equivalent to physiology : recently, it has come to denote the natural history of plants and animals, i. e. of the two organic kingdoms, including both their physiology and descriptive natural history. 1 2 INTRODUCTION. of plants and animals are composed, is called organic matter. Animal- appropriate ami live upon this, but have not the power of producing it. So the \egetablc kingdom stands between the mineral and the animal ; and its function i> to convert materials of the one into food for the other. Although plants alone are eapalile of building up living structure out of mineral mate- rial-, and are the sole producers of the organic matter which U c--ential to animal life, and although animals consume that which plants produce, vet plants also consume organic matter. more or less, acting in this respect like animals in all their opera- tions, except in the grand and peculiar one by which they tissiniiltiti- mineral matter. Most plants of the higher grades a»imilate largely and consume little, except in special opera- tions. Sonic, on the contrary, are mainly consumers, and feed upon formed organic matter, living in this respect after the manner of animals. The living substance of plants and animals is essentially the same. 4. Botany deals with plants : 1. As individuals, and in respect to their structure and functions. 2. In their kinds, and as respects their classification, nomenclature. &c. Accordingly, the most comprehensive division of the science is into Pmsio- LOGICAL or BIOLOGICAL BOTANY (using these terms in their widest sense) and Svsi I.MATIC BOTANY. But as Physiology and Biology, in the restricted sense, relate only to functions or actions and their consequences, the lir>t department naturally divides into two, \\y.. Structural Botany and Physiology. .".. SIKM KUAL BOTANY comprehends all inquiries into the structure, the parts, and the organic composition of vegetables. This is termed OKGANOGKAIMIY, when it considers the organs or obvious parts of which plants are made up, and MORPHOLOGY, when the study proceeds on the idea of type. The term < >ia, \\oi.i NY lias been applied to the study of the nascent organs and their development; PIIYTOTOMY, or VK<;KTABLE ANMOMY. to that of the minute structure of vegetables as re- vealed by the microscope, i. e. to the composition of the organs themselves, lint, since anatomy in the animal kingdom includes the consideration of general as well as of minute structure, and indeed answers to organography, the minute anatomy of both kingdoms takes the special name of HISTOLOGY. The study of functions, or of the living being (animal or plant) in action, is the province of I'm SIOI.OCY. i1'. SYSTKMATK I'.OCVM. or the study of plants in their kinds and in regard to their relationships, comprises TAXONOMY, or the principles of classification, as derived from the facts and ideas INTRODUCTION. 3 upon which species, genera, &c., rest; CLASSIFICATION or the SYSTEM OF PLANTS, the actual arrangement of known plants in s}Tstematic order according to their relationships ; PHYTOGRAPHY, the rules and methods of describing plants ; and NOMENCLATURE, the methods and rules adopted for the formation of botanical naines. GLOSSOLOGY or TERMINOLOGY 1 is a necessaiy part of Phytography or Descriptive Botany, and hardly less so of Structural Botany : it relates to the application of distinctive terms or names to the several organs or parts of plants, and to their numberless modifications of form, &c. This requires a copious vocabulary of well-defined technical terms, by the use of which the botanist is able to describe the objects of his study with a precision and brevity not otherwise attainable. It will be convenient to exemplify the principal terms along with the modifications of conformation which they designate ; and also, for greater fulness and facility of reference, to append to this volume an alphabetical summary of them, or Vocabulary of Botanical Terms.2 7. The present volume is mainly devoted to Morphological Botany ; that is, to Structural Botany on the basis of mor- phology. This department cannot be properly dealt with apart from considerable reference to intimate structure, development, and function, the subject-matter of vegetable histology and physiolog}'. But these will here be treated only in the most general or incidental and elementary wa}' , and only so far as is necessary to the understanding of the morphology of the stem, leaves, &c. The whole discussion of the histology and physiology of plants is relegated to a following volume and to another hand. 8. The most comprehensive and important division of the vegetable kingdom is into plants of the higher and of the lower series or grade, i.e. into PH^ENOGAMOUS (or PHANEROGAMOUS) or FLOWERING, and CRYPTOGAMOUS or FLOWERLESS PLANTS. The first are all manifestly of one t}^, and therefore have a consist- ent and simple morphology. The second differ among them- selves almost as widely as they do from the higher series ; and 1 GLOSSOLOGY is the better word, but TERMINOLOGY, although a hybrid of Latin and Greek, is in common use. 2 What is called GEOGRAPHICAL BOTANY is the study of plants in respect to their natural distribution at the present time over the earth's surface, and the causes of it. FOSSIL BOTANY (Vegetable Palaeontology) relates to the plants of former ages, as more or less made known in their fossil remains. MEDICAL BOTANY, AGRICULTURAL BOTANY, and the like, are applications of Botany to medicine, agriculture, &c. 4 INTRODUCTION. their morphology is more special and difficult. Wherefore it is better to treat them separately and subsequently. This will be done in a third part, by an associate devoted to Cryptogamic Botany. 9. Thus the field is here left clear for the Structural Botany of Phaenogamous or Flowering Plants, with which the study of the science should naturally begin. In theory it may seem proper to commence with the simplest plants and the most ele- mentary structures ; but that is to put the difficult and recondite before the plain and obvious. The type or plan of the vegetable kingdom, upon which morphological botany is grounded, is fully exemplified only in the higher grade of plants, is manifest to simple observation, and should be clearly apprehended at the outset. CHAPTER I. OUTLINES OF THE GENERAL MORPHOLOGY OF PH^ENOGAMOUS PLANTS. 10. MORPHOLOGY, the doctrine of forms, as the name denotes, is used in natural history in nearly the same sense as the older term Comparative Anatomy. If it were concerned merely with the description and classification of shapes and modifications, it would amount to little more than glossology and organography. But it deals with these from a peculiar point of view, and under the idea of unity of plan or type.1 11. As all vertebrate animals are constructed upon one type (or ground plan), which culminates or has its archetype in man, so all plants of the higher grade (8) are strictly of one type ; the different kinds being patterns or repetitions of it, with varia- tions. The vegetable kingdom, however, does not culminate in an archetype or highest representative. As respects the organs of vegetation, the higher classes of cryptogamous plants exhibit this same type ; but it is only in the most general or in a recondite sense that this can be said of their organs of repro- duction, and of the less differentiated structure of the lowest classes. Wherefore cryptogamous plants are left out of the present view, to be treated apart. 12. Viewed morphologically and as to its component organs, a plant is seen to consist of an axis or stem, which sends off roots into the soil, and bears lateral appendages, commonly as leaves, but which may be very unlike leaves in whole appearance 1 The term Morphology was introduced into science by Goethe, at least as early as the year 1817 (Zur Naturwissenschaft iiberhaupt, besonders zur Morphologie, Stuttgart und Tubingen, 1817-24). On page 9 of the first volume, he is understood to have suggested this word for the purpose and in the sense now adopted in botany and zoology. It essentially replaces an earlier and somewhat misleading word, Metamorphosis. (304.) Apparently the first botanist to adopt the term was Auguste de St. Hilaire, in his " Le9ons de Botanique, comprenant principalement la Mor- phologie Ve'ge'tale, etc., Paris, 1841. The term seems not to have been taken rup, in zoology, by Etienne Geoffrey Saint-Hilaire, the antagonist of Cuvier (who was of a wholly different family from that of the botanist), although the same idea was denoted by his phrase " unity of organic composition " 6 GENERAL MORPHOLOGY :me< -n 1>V botanists extendeil iii a generic way from the green expan- sion- which constitute foliage to other forms under which such appendages occur. The proper morphological e\pre--ion is. that the latter are homologous with leaves, or are the Ao//>oA/y"< > of leaves.1 13. Leaves are borne upon 'the stern at definite places, which are termed NODES. A node may bear a single leaf or a greater number. When it bears two, they occupy opposite sides of the stem. When three, four, or more, they divide tin- circum- ference of the stem equally, forming a circle, technically a WHORL, or in Latin form a VERTICIL. When only two. the pair evidently answers to the simplest kind of whorl. So that leaves are either single on the nodes, in which case they arc alter- nate, that is, come one after another on the stem : or in whorls (ichorled, verticillate) , in the commoner case of a single pair being called opposite . The bare space between two successive nodes is an INTERNODE. This is longer or shorter, according to the amount of longitudinal growth, which thus spaces the leaves, or whorls of leqves, in most various degrees, either widely when the internodes are elongated, or slightly when they remain very short. The plant, therefore (roots excepted), is made up of a series of similar parts. /'. c. of portions of stem, definitely bearing leaves, each portion developed from the apex of the preceding one. This constitutes a simple-stemmed plant. 14. Branching is the production of new stems from the older or parent stem. These normally appear in the AXILS of leaves, that is, in the upper angle which the leaf forms with the stem. - from which they grow much as the primary stem grew from the seed. The primary stein, connected with the ground, produco roots which develop downwardly into the soil, from which they draw sustenance. Branches, when developed above ground. 1 A nmiintm designation for all these appendages bciiiL; desirable. :i £ 1 one is 1'imiisheil by tin- (ireek name for leaf, n>/>hijllous, hypsophyttous, gamophyllous, &c. OF PH^ENOGAMOUS PLANTS. being in organic connection with their parent stem, do not usually produce roots ; but when placed in equally favorable conditions for it, i. e. on or in the soil, they may strike root as freely as does the original stem. 15. An incipient stem or branch, with its rudimentary leaves, is a BUD. The normal situation of a bud is in the axil of a leaf (axillary} , the development giving rise to branches ; or else at the apex of an axis (terminal), where there can be only one, the development of which continues that axis.1 16. As branches are repetitions and in one sense progeiry of the stern which bears them, so the serial similar parts or leaf- bearing portions of a simple stern are repeti- tions, or in a like sense progeny, each of the preceding one from which it grew. The simple-stemmed plant is made up of a series of such growths, each from the summit of its predecessor ; the branched plant, of ad- ditional series, laterally developed, from ax- illary buds. These ultimate similar parts into which a plant may thus be analyzed, and which are endowed with or may produce all the fundamental organs of vegetation, were by Gaudichaud called PHYTONS. But phyton, being the common Greek name for plant, was not a happily chosen appellation for plant- elements, or homologous plant-units. A better term for them is PHYTOMERA (cpvrov, plant, ftsQo^, part), equivalent to plant-parts, --the structures which, produced in a series, make up a plant of the higher grade. In English, the singular may be shortened to PHYTOMER. 17. This theoretical conception of the organic composition of the plant is practically impor- tant to the correct understanding of morpho- logical botany. The diagram, Fig. 1, serves to represent the organic elements, or phi/fomera, in a simple case, such as that of a growing plant of Indian Corn, or other Grass. Here 1 Bifurcation by the division of a terminal bud into two, as in Acrogenous Cryptogams, is supposed by some to occur, even normally, in some Plueno- gams, especially in certain forms of inflorescence ; but this has never been convincingly made out. FIG. 1. Diagram of a simple-stemmed plant, exhibiting the similar parts, or phytomera, a to h, of which it is composed. 8 GENERAL MORPHOLOGY the leaves are alternate; in other words, each phytomer is single-leaved ; while in the subsequent illustrations of plants developed from the seed, at least the earliest phytomera are two-leaved. 18. The plan thus exhibited in the leafy stem begins in the embryo, or initial plant in the seed, and is carried on into the flower, in whieh the normal development of the axis iinally ends. One plan prevails throughout. To illustrate it, the morphology and growth of the embryo, of the plant developed for vegetation and the general purposes of its individual existence, and lastly of the flower, through whieh sexual reproduction takes place, may be successively treated in this order. OF THE EMBKYO AND SEEDLING. CHAPTER II. MORPHOLOGY AND DEVELOPMENT OF THE EMBRYO AND SEEDLING. 11). The Embryo is the initial plant, originated in the seed.1 In some seeds it is so simple and rudimentary as to have no visible distinction of parts : in others, these parts may have assumed forms which disguise their proper character. But every well-developed embryo essentially consists of a nascent axis, or stem, bearing at one end a nascent leaf or leaves, or what an- swers to these, while from the other and naked end a root is normally to be produced. This stem is the primitive internode of the plant : its leaf or pair of leaves is that of the first node. The plant therefore begins as a single phytomer. Some embryos are no more than this, even when they have completed their proper germination : others have taken a further development in the seed itself, and exhibit the rudiments of one or more fol- lowing phytomera. The embryo of the Maple is an example of the first kind ; and, being large enough for handling and for the display of all its parts to the naked eye, and the character of these parts being manifest even in the seed, it is a good subject with which to commence this study. And for this the Sugar- Maple is one of the best of the Maples. Its embryo (seen in Fig. 2 in the coiled condition which it occupies in the seed, and in Fig. 3 and Fig. 4 uncoiling and be- ginning to grow) is an initial stem, bearing a pair of leaves, and nothing more. These parts take the technical names of 1 Normally a seed contains a single embryo. Polyembry, the formation of two or more embryos, occurs occasionally as a kind of superfo3tation in some seeds. In those of the cultivated Orange it is most common, and an evident monstrosity. In Coniferae and Loranthacese, two or three embryos, of equal size and perfection, are not rarely produced. FIG. 2. Embryo of Sugar Maple, in vertical section, as coiled in the seed, merely Bomewhat loosened. 3. Embryo of same, just beginning to unfold in germination. 4. Same more advanced : a. its stem or caulicle ; «». its two leaves or cotyledons. 10 MORPHOLOGY 20. Caulicle or Radicle, and Cotyledons. The name of radicle was early applied to tin- uxis of tin- embryo below the cotyledons, on the supposition that it was the actual beginning of the root. But its structure and mode of growth show it is not root (iM. 11. 7*), but a body of the exact nature of Mem. from the naked end of which the root is developed. Wherefore ('iuiln.lt' (Lat. canllcitluK, diminutive of mulis. stem) is tin- appropriate name : and it would he gen- erally adopted, wen' it not that the older term is so incorporated into the language of sys- tematic 1 iot any (in which fixity and uniformity are of the utmost importance) that it is not easily displaced. It may be continued in descriptive botany on this account, but in morphology it is apt to mislead ; and the name of caulicle, suggestive of the true nature of the organ, is preferable.1 The more fanciful name of Cotyledons was very early applied to what are now recognized as answering to the leaves of the embryo : it has the negative merit of suggesting no misleading analogy. '- 21. Development of the Dicotyledonous Em- bryo, /. e. the two-leaved embryo. This, in the Red Maple (Figs. 5-8), usually germinates in summer, shortly after the fruits of the season have matured and fallen to the ground. It differs from that of Sugar Maple in the crump- ling instead of coiling of the cotyledons in lin- seed. Referring the whole physiology of ger- mination to that part of the work which treats Of Vegetable Physiology . the development of the embryo into the seedling may hen- be described, taking that of a .Maple fora convenient type or pattern, with which other forms 1 Linnaeus called it Hoodlum, a nanu- which, bein"; etymological] v mean- ingless in this connection, is not misleading. The French lioiani.-ts named it Tii/ill'-, diminutive of tiI/. Seed extrnrled ami divided, to show the embryo within. 7. Embryo parti*1 vuil'oldiMl. 8. Embryo in early sta^e of OF THE EMBRYO AND SEEDLING. 11 ma}' afterward be compared. The first growth is seen in the elongation of the radicle or caulicle, and its assumption, as far as possible, of a vertical position, and the production of a root from the naked end. As it emerges from the seed in consequence of this elongation, the root-end of the caulicle points downward into the soil, the caulicle bending, if need be, to assume this position ; and the nascent root, partaking of this disposition, grows in a downward direction. Hence the root has been called the Descend- ing Axis of the plant. While this avoids, the opposite or budding end (as it may be termed) seeks the light, and when free takes an upward direction. The result of this, and of the elongation of the caulicle, is to carry the budding end out of the soil and into the air, where the growing cotyledons unfold or expand and become the first leaves, or Seed-leaves. This initial stem and its continuation therefore constitutes the Ascending Axis. If the budding end happen to lie pointing downward and the root-end upward in the ground when germination begins, both will curve quite round, as the}' grow, to assume their appropriate directions. If obstacles intervene, each will take as nearly as possible its wonted direction, through an instinctive tendency and action, which insures that each part of the plant shall be developed in its fit medium, — the root in the dark and moist earth, the stem and leaves in the light and air. 22. The plantlet, thus established, has now all the essential Organs of Vegetation, as the}' 'are called, i. e. root, stem, and leaves. Its subse- quent development, so far as vegetation (apart from proper reproduction) is con- cerned, consists in the addition of more of these, until the whole herb, shrub, or tree is built up. 23. In Maples (as in the Morning Glory, Fig. 16, and many others) the embryo in the seed, and until after the full develop- ment of its cotyledons or seed-leaves, shows no rudiments of the subsequent growth. The embryo grows into the plant- let wholly by the appropriation of prepared nourishing matter which was provided by the mother-plant and stored in the seed, — in the case of the Maple, wholly in the embryo itself, mainly in its cotyledons. FIG. 9. Maple embryo developed into plantlet of one phytomer, and producing rudiments of the second: the lower portion covered with root-'wjrs is the root; tin* naked portion above is the caulicle. 12 -MORPHOLOGY After this is consumed and in good part converted into struc- ture, the plantlet must l>y the action ol' its root and leaves imbibe from the soil and air appropriate materials, and assimilate them into nourishing matter needful for further growth. Only then does the rudi- ment of new structure appear, in the form of a growing point, or bud. at the node or apex of the primitive stemlet. between the two seed- leaves. In this case it soon .shows itself as a second pair of leaves, at first resting on the node (Fig. 'J). next as somewhat upraised by the development of the second intcrnocle (Fig. 10, summit), and finally both this inter- nodi1 and the pair of leaves complete their growth (Fig. 11). Then the terminal bud which crowns the second node develops in the same way the third pair of leaves and their supporting inteniode or joint of stem (Fig. 12) ; and so on. 24. The root and the stem grow not only in opposite directions, but in a different mode. The primordial stem, pre-existing in the seed (though at first it may be extremely short) grows throughout its whole length, but most in its upper part, so that it may become a stemlet two or three inches long. l>ut. soon attaining its full growth as to length, the stem is carried upwards by the subsequent joints or portions, similarly developed and elongated, one after the other. Not that each portion necessarily waits until the growth of its prede- cessor is complete, — though this occurs at first in seedling Maples and other embryos unprovided with much store of food, — yet the development follows this course and order of succession. The root, on the contrary, cannot be said to pre-exist in the seed, or at most it may be said to exist potentially in tissue of the caulicle from which a root or roots normally originate.1 It is formed 1 Vet from nothing which is special to this part of tin- embryo, nor to the embryo at all. The primary root is developed from subjacent tissue of the tip of the eauliele. just MS it is sometimes developed from ahmu the sides, and as secondary roots are from all or must Mems under favoring conditions. This complete similarity, and the fact of \vhat is called the "endogenous" origin of roots (i.e. their sprinirinir from subjacent rather than superficial tissue) appear fully to warrant the statement in the text above. Fit!. 10. Mapli1 plantlet with scniml internode developing. 1). Same with second nair of li>;ivr- rimijili-ti1. anil bud of the third apjiavrnt. OF THE EMBRYO AND SEEDLING. in the process of germination, and originates in tissue just back of that which covers the root-end of the caulicle, and which, being carried forward by the subjacent formation (to which it becomes a sort of cap or sheath) , is called the Root-cap. As the primary root thus began by a new and local growth at the extremity of pre-existing stem, so it goes on to grow in length wholly or mainly by a continuation of this formation, the new at the end of the old. That is, the root elongates by continual minute increment of its apex or near it, the formed parts very soon ceasing to lengthen. This is in marked distinc- tion from stem, which grows by suc- cessive individualized portions ; and these portions (internodes) , at first very short, attain or are capable of attaining a considerable and sometimes very great, but definitely terminable length, by interstitial growth through- out. Moreover, roots are naked, not producing as the}' grow either leaves or any organs homologous with leaves. The}' commonly branch or divide, but in a vague manner ; and their new parts bear what are called Root-hairs, which greatly increase the absorbing surface ; other- wise they are destitute of appendages or organs. 25. With the Maple embryo, here taken as a type, that of Morning Glory, Ipomoea purpurea, or an}7 of its kin, may next be compared. The cotyledons are different in shape, being as broad as long, and notched both at base and apex. They lie in contact in Fig. 14, and are very thin, leaf-like, and green while contained in the seed. Their thinness is shown in Fig. 13, where a section of the crumpled and folded embryo, as it lies in the seed, exactly divides them (passing through the terminal and basal notches) and also the caulicle, which here is thicker than both. The germination is similar to that of the Maple ; and like that (as Fig. 16 shows), and for the same reason, no bud or rudiment of the further growth pre-exists in the embryo or FIG. 12. Red Maple seedling, with three joints of stem and pairs of leaves developed, the first being the cotyledons. 14 MORPHOLOGY appears in the young plant let, until that has established itself and hail time to elaborate proper material therefor. This con- dition U correlated with thin foliaeeous cotyledons, holding no store of noiiri-h- ineiil. Here they do not contain sullicieiit material for the development of the initial stem and root. The maternal provision for this is here stored up in the seed around hut not within the embryo. This nourishing deposit, seen in the section (Fit;-. \:\) filling the whole space iietween the seed-coats and the thin embryo, was named by the early botanists and vege- table anatomists the Ai.r.t MKN of the seed.1 This substance, softened in germination and by chemical changes rendered soluble, is gradually absorbed by the cotyledon^ as material for their growth and that of the developing primary stem and root. 26. Seeds in this regard are accordingly distinguished into albuminous and <>.nil- l>itin!>i/«/•/«, and Richard by /•.'//•/»/•///, neither of them much better etyinolo^ically than the old word Allmnn n. But it must be kept in mind that it was intended to liken the " albumen " of the seed with the albumen or white of an CLIU' as a liody or mass, and not as a chemical suh-taiiee; the embryo bein.u' fancifully conceived to be analo- gous to the //'nnca purimrea. ui- spi cad as the tirsl pair of leaves. 16 OF THE EMBRYO AND SEEDLING. 15 portional to the size and strength of the embiyo, or the degree of its development in the seed. A comparison of the various illustrations sufficiently shows this. Figures 17 to 24 exhibit, in a few common seeds, somewhat of this relation, and also of the position and shape assumed in some instances. The upper rank of figures represents sections of seeds ; the embiyo left in white ; the albumen as a dotted surface. The lower rank shows the embryos detached. That of Mirabilis has very broad and thin cotyledons, a caulicle of equal length, and the whole curved round the albumen which thus occupies the centre of the seed. That of Potato is coiled in the midst of the albumen, is slender ; the cotyledons narrowed down to semi-cylindrical bodies, not leaf-like in appearance, and the two together not thicker than the caulicle. In Barberry the embryo is straight, in the axis of the albumen, which it almost equals ir. length ; the cotyledons considerably broader' than the caulicle, but short and thickish. That of the Peony is similar, but very much smaller, occupying a small space at one end of the albu- men, and seemingly without distinction of parts, but under the microscope and with some manipulation the broader end is found to be divided, that is, to consist of two minute cotyledons. The embryo of a Crowfoot is similar, but still more minute and the parts hardly to be distinguished ; and in some minute em- biyos there is no apparent distinction of parts until they develop in germination. 28. The study of the formation of the embiyo in the seed teaches that all embryos begin with a still more simple, minute, and homogeneous structure ; and these comparisons suffice to show that all such differences are referable to different degrees and somewhat different modes of the development of the embiyo while yet in the seed. It also appears that the size and shape FIG. 17. Section of seerl and contained embryo of Mirabilis (Four-o-clock). 18. Embryo detached entire. FIG. 19. Section of a Potato-seed. 20. Embryo detached entire. FIG. 21. Section of Barberry-seed. 22. Embryo detached entire. FIG. 23. Section of Peony-seed. 24. Embryo detached entire. 16 MORPHOLOGY of an or-ian do not indicate its nature, either in the embryo or in subsequent growth. But in :dl the cases yet mentioned the cotyledons actually demonstrate their nature by developing in germination in a foliaceous manner and becoming the first leaves of the seedling. Nor is this nature much disguised by the fact that they differ greatly in form in dillerent species, and that the seed- leaves, or developed cotyledons, differ much in shape and often in texture from the succeeding leaves. ( See Fig. 11, 12, 25, &c.) 29. To complete the comparison between the seedling Morning Glory and that of the Maple, it is to be noted that here, while the cotyledons or seed-leaves are two, the following internode bears only one leaf (Fig. 25), as also will the just de- veloping third internode ; and this continues throughout up to the blossom : that is, the leaves subsequent to the cotyledons are not opposite as in the Maple, but alternate. (13.) 30. All the preceding illustrations are from embryos which previous to germination have developed nothing beyond the cotyledons. In the following, a rudiment of further growth, 25 26 27 28 29 30 or a primary terminal bud, is visible in the seed. It is most manifest in large and strong embryos with thick or fleshy cotyle- Fltl. 'jr.. Further development nf Morning , the root cut away, the iiiterin.de :iliovc the cotyledons Mini its leaf completed, I lie next intcrnode ;inil its leaf appearing. FI<;. •_'(•,. Embryo (kernel) of the Almond. 27. Same, with one cotyledon removed, to show I he plumule, a. FIG. 28. Section of an Apple-seed, magnified, cutting throiiu'h the thickness of the cotyledons I'll Kmhr\ o of the same', ex I rue ted entire, the cotyledons a little separated. FKi. :til. (ieniiiiiali in of the ( 'herry, showing the thick cotyledons little altered, and the plumule de\el >pin:,' the' earliest real foliage. OF THE EMBRYO AND SEEDLLMJ. 17 dons, i. e. cotyledons well charged with nourishing matter. The early vegetable physiologists gave to it the name of PLUMULE (Lat. plumula, a little plume). The name was suggested by its appearance in such an embryo as that of the bean (Phaseolus), in which it evidently con- sists of a rudimentary pair of leaves, while in the pea and the acorn it is a rudimentary stem, the leaves of which appear only later, when germination has considerably advanced. In any case, the plumule is the bud of the ascending axis already discernible in the seed. Fig. 27, a, shows it in the almond, one cotyledon being removed. Fig. 28 shows it in the section of a similar although much smaller embryo, that of an apple-seed, enlarged to nearly the size of the other. It is equally visible in the cheriy, the bean, and the beechnut. The embryo in all these cases constitutes the whole kernel of the seed. For the nourishment, which in all the foregoing illustrations except the first (i.e. in Fig. 13, 17-23), is deposited around or exterior to the embryo, is in these stored within it. 31. The development of these em- biyos in germination proceeds in the normal manner, but with two cor- related peculiarities. First, by the lengthening of the radicle more or less, their thick cotyledons are usually raised to or above the surface of the soil ; the}- expand, assume the green color needful to foliage ; but they imperfectly or in a small degree perform the function of green leaves. Their main office is to supply the other growing parts with the prepared nourishment which they abundantly contain. Then, being thus copiously nourished, the root below and the ready-formed plumule above grow rapidly and strongly, having accumulated capital to draw upon ; and the leaves of the FIG. 31. Beechnut cut across, filled by the fleshy embryo; the thick cotyledons partly enfolding each other. 32. Embryo of the same in early germination 33. Same more advanced ; the plumule, which is just emerging in the preceding, here developed into a long iiiternode and a pair of leaves. 2 18 MORPHOLOGY latter arc practically the earliest cllieient foliage of the plnntlet. Thus, as in the germinating C'lierry-seed ( l-'i^-. :;i»). three or four internodes of stem, with their leaves, may be produced before these leaves themselves are siillicieiitlv developed to make any contribution to this irrowth. And in the Beech :,nd lieaii. the leaves of the plumule come forward almost before the root has attached the plantlct to the soil. (FiiT. -'^. :i;").) Between such cases and that of Maple and the like there are all decree >. There are also familiar east's in which the storage of nourishment in the cotyledons is carried to a maximum, with results which gravely ailed the development. j.'Ki. :;i. '['!,,. rnil.r\n Mhr \\lml,' krnn'l) of the Bean. 35. Same early in ^ermi- TKition; (lie tliiiU e.iiyleilniis e\|i:»ncliii'4 ami slinwiii^ the plumule. 3G. Same, more julvanei'il in uenninatinii ; (lie plumule Jovelxpeil into :ui iiiterncule nf stem liearing a ]i:iir 1. 1' le:i\ 88. FIG. :!7. Kinbryo of Pea, i. <•. a pea minus the seed-coat. 38. Advanced germi- natior of the same OF THE EMBRYO AND SEEDLING. 19 32. Thus, in the Pea, near relative of the Bean, the embryo (Fig. 37), which is the whole kernel of the seed, has the cotyledons so gorged with this nutritive store that they are hemispherical ; and the acorn of the Oak (Fig. 39), near ivlative of the Beech, is in similar case. These extremely obese cotyledons have not only lost all likeness to leaves, but all power of fulfilling the office of foliage, which is apparently no disadvantage ; for when two different duties are performed by the same organ, it rarely performs both equally well. Here they become mere receptacles of prepared food, the nature and office of which is the same as of the albumen, or nutritive deposit exterior to the embryo in what are called albuminous seeds. (25-27.) The difference is in the place rather than in the character of the deposit. The plumule in such cases is always apparent before germination ; and it develops even with more vigor than in the preceding cases. It usually rises as a stout stem of several internodes lengthen- ing almost simultaneously, or at least the upper strongly developing long before the lower have finished their growth ; and the latter are practically leafless, bearing only small and scale-like and useless ru- diments of leaves. This is correlated with the peculiarity that the caulicle does not lengthen in germination, or it lengthens very slightly ; the cotyledons remain within the coats of the seed ; and if this were buried beneath the surface of the ground, there it remains. The abortion of the earliest leaves of the plumule is in correlation with this hypogaous (i. e. underground) situation of the cotyle- dons throughout the germination. The slight elongation of the caulicle serves merely to protrude its root-end from the coats of the seed in a downward direction, and from this a strong root usually is formed. FIG. 39. Section of an acorn, filled by the embryo. 40. Advanced germination 01 the same. MORPHOLOGY 33. In some Oaks, notably in our Live Oak (Quercus virens), ainl less so in tin- Horsechestiiut. the two cotyledons coalesce or cohere by their contiguous faces. In sonic <>f these C.-IM-S nl' hypo- irii-ous germination. the short caulide .•Hid plumule arc extri cated from the enclosing coats or husk by the development of short stalks (petioles, l.'.Ti to the fleshy cotyledons : as is seen in Fig. 42, and in most germinalin-- acorns. These petioles arc not visible in the seed, but arc the first develop- ment in germination. 34. There are some curious cases in which, while the caulicle remains short and subterranean. the cotyledons are raised out of ground in germination by the format ion of far longer stalks (petioles) than those of the Horsechestiiut. A singularly dis- guised instance of this kind is seen in Megarrhiza, a genus of Cucurbitaceoiis plants of California and Oregon, remarkable for their huge root. The large seed has very thick and fleshy cotyledons, and a very short and straight caulicle. In germi- nation, the whole seed is elevated, seemingly in the manner of the bean, upon a stout stem. One waits for a long time expect- ing to see the cotyledons throw oil' the bursting husk and expand, or else to put forth the plumule from between their bases. But at length the plumule, makes its appearance from an unexpected place, coming separately out of the soil. Removing this. (In- state of things represented in Fig. 1:5 is presented, -that of the plumule seemingly originating from the base, instead of the apex, of an elongated caulicle ! But on examination of the cleft from which this proceeds, by making a section of the stem above (showing that it is hollow), and finally by separating the cotyle- dons and gently tearing apart the two short stalks by which they are united to their stem-like support, it is found that the latter may be divided into two (as shown in Fig. 44). even down to the deft below. This explains the anomaly. The real caulide has re- Kic 41. Section of M Horsrrhrstnnt or I'.uH-rvr srr.l. tlironrrh the very thirk rotylnlons and I lie incurv.'il ciinlicle. 42. Seed in perminntion. showing the peti to tin' rotylrdiins, >V < OF THE EMBRYO AND SEEDLING. 21 mained short and subterranean, and is confluent with the upper part of the thickening root : the seeming caulicle, which raised the cotyledons above the soil, consists of the petioles of these combined into a tubular stem-like body, no evident trace of which is visible in the seed, although in germination it attains the length of two or three inches : in age it is readily separable into the two leaf-stalks or petioles of which it is composed : the plumule is thus seen to be wholly normal, originating from between the cotyledons. All the ex- tensive growth so far, and until the proper foliage-leaves of the continu- ation of the plumule are developed and begin their action, is from nutri- tive material stored in the thickened cotyledons, a considerable part of which was transferred to the already enlarging root, before a remaining portion was used in building up the strong plumule . The economy of this elevation of cotyle- dons which never open, and of the lengthened distance through which the nutritive matter has to be carried, is not apparent. But it is the family habit in Cueurbitaceae to bring up the cot^yle- dons that they may develop as leaves (as in the Pumpkin, Fig. 47) : here this elevation is brought about in a different wa}', but without securing the useful end. l 1 It may be inferred that Megarrhiza is a descendant of some Cucurbitacea with thinner cotyledons, which in germination developed into long-stalked leaves, in the manner described in the next following paragraphs. FIG. 43, 44. Peculiar germination of Megarrhiza Californica; explained above. -2-2 MORPHOLOGY 35. This same anomaly, as to the development of long stalks to the cotyledons and their union into a stem-like body, occurs in various ^ici-ie- of Larkspur (notably in the California!) Delphin- ium nudicaule) : but in these the cotyledons develop into a pair of ellicicnt liTcen lea\ 68. 45 47 3G. A similar elongation of petioles of the cotyledons, but without any union, occurs in a species of Morning (ilory of the plains beyond the Mississippi (Ipoma-a leptopliylla) : the leaf- like cotyledons coming up on their long stalks separately from the ground (Fig. 45) ; the developed plumule rising some time afterward between them. Compare this with the ordinary species (-J."). Fig. 1,"). 1C. -_'.".). and note that the dill'erence is merely that the caulicle in the common Morning ( < lory elongates and the petioles of the cotyledons remain short. 37. In all instances thus far a single primary root so regularly develops from the lower end of the axis of the embryo (variously named radicle or cauliele) . and forms such a direct downward !•'!<;. )."•. Germination of I|M>HIM-:I leptopliylla; the caulicle not developing, the plumule and the pi-tiolnl rotylnlons rise from unilercrouiul. Dotted line marks the level of tlie soil. Kir,, ic. Embryo of a Pumpkin, the cotyledons separated. 47. Same germinated; a cluster of routs from tin- l>:isc of caulicle. OF THE EMBRYO AND SEEDLING. 23 prolongation of it, that it was called the descending axis ; and the bod}' from which it originates was named the radicle, on the supposition that it was itself the nascent root. But, as already explained, the so-called radicle grows in the manner of stem (24), and is morphologically that initial internode the node of which bears the first leaves or cotyledons. (20.) Let it now be noted that this descending axis or single primary root is far from universal. In Pumpkin, Squash, Echinocystis, and the like, the strong caulicle sends out directly from its root-end a cluster of roots or rootlets, of equal strength; i. e., it strikes root in nearly the manner that a cutting does. (Fig. 47.) 38. The Polycotyledonous Embryo is one having a whorl of more than two seed-leaves. The dicotyledonous embryo being a whorl of the very simplest kind, that is, with the members reduced to two, the polycotj-ledonous may be regarded as a variation of it. In all but one group of plants it is simply a variation, of casual occurrence, or even a monstrosity, in which three or rarely four cotyledons appear instead of two. In Pines (Fig. 48, 49), however, and in most but not all Coniferse, a whorl of from 3 to 10 cotyledons is the normal structure, varying accord- ing to the species, but of almost uniform number in each. In germination these are brought out of the soil by the elongation of the caulicle, and when the husk of the seed is thrown off they expand into a circle of needle-shaped leaves. In the Pine tribe, all the subsequent leaves are alternate (spiral) in arrange- ment, with some disguises. In the Cypress tribe, the cotyledons are fewer (not more than four, and more commonly only two), and the subsequent leaves also are in whorls of two to four ; i. e., are either opposite or verticillate. From the occasional union at base of the cotyledons of a polycotyledonous embiyo in pairs or groups, and from a study of their early development. Duchartre l plausibly maintains that such cotyledons really consist of a single pair, parted into divisions or lobes. The ordinaiy interpretation, however, is equally tenable. 39. The Monocotyledonous Embryo, although theoretically the simplest, is practically a more difficult study. It has a single cotyledon (as the name denotes) ; also a single leaf to each node 1 Ann. Sci. Nat. ser. 3, x. 207. This view, which originated with Jussieu, is adopted by Parlatore in DC. Prodr. xvi. FIG. 48. Section of a seed of a Pine, with its embryo of several cotyledons. 49. Early seedling Pine, with its stemlet, displaying its six seed-leaves. -MoUi-ilOLOGY 54 of tlir plumule • thai is, the leaves of the embrvo are alternate, lint the eauliele is usually very short, ami there i> u,. external niark by which its limiN may l.e di>tin- guUhed from the cotyledon, until gcrmi- nation ha- begun. For a type of it. the embryo of some aquatic or mar>h plants may lie taken. \vh. iv it forms ihe whole k''1'1"'1 of the seed (Fig. 50-53), and 63 the structure can he made out antecedent to germination. It is understood l.y supposing that the cotyle- don, which forms its principal bulk (the caulicle being only the very short thickish base), is convolute around a short plumule, and the margins concreted, except a minute longitudinal chink at base, out of which the growing plumule protrudes in germination. The embryo of his may he similar in structure, hut no distinction of parts is visihle. It is very small in proportion to the size of the seed, the kernel being mostly albu- men, — a supply of food, from which the germinating embryo draws the materials of its growth. When this takes place, either the cotyledon or the whole embryo lengthens, its lower part is pushed out of the seed, a root forms at the free end of the excessively short caulicle. and the plumule develops from the other in a series of one-leaved nodes, the internodes of which remain so short that the leaves continue in close contact, the bases of the older successhely enclosing the inner and younger. (Fig. ,">5.) Here. therefore, the cotyledon mainly remains in the seed, and the seed remains underground ( hypog;eoiis) . 40. It is somewhat dilferent in the Onion, which has a similar embryo, except that it is longer, and the cotyledon is curved in the albumen of the seed. The iirsl steps are 1 he same as in Iris; but as soon as a root is formed and embedded in the soil, the cotyledon lengthens vastly more, into a long and liliform green leaf, which, taking an erect position. I'll 1 r.n. S 1 ofTri.nlcirliiii |i.-ilnsliv; I lie rh.-ij.hr. Ir.'iilinj; In tin • str. •!!;,' clialaxa :it tlir Miiinnil. tiiru-'il tnwiirils I In •<•><• 51. Tl mlu-> •,> ilrt.-irlicd IVuni tbe Seed-COatS, ^Imwiiig the longitudinal chink at the base of the cotyledon; tlie short part below ia the radicle. \vi Hi tin' i-li ink timinl l:itcr:illy, and half the cotyledon CUl awny. IPI infill:,' to \i.'\\ Hi.' |iliniiuli' i-oiiccMli'd within. '•'.'.. A rr,iss-srr| i,m through the plunml.-, inure •ilicil. ]•'!! ;. .". I. Scclioii .if. -ci'il nf Iris. I'lil.ir.,'.'.!. >.li.i« IIIL; tin- sin rill ninl .-iiiiciri-ntly siinplr c!iiiir\i> at the base of the albumen .r>.".. Ccnninatim: sr.'.i nn.l ser.liiir,' ,.i tin- s; ..f nntiinil si/c. OF THE EMBRYO AND SEEDLING. 26 carries up the light seed far above the surface of the ground, the tip only remaining in the albumen of the seed until that is ex- hausted, when the tip perishes and the emptied husk falls away. About this time the plumule shoots forth from one side of the subterranean base of this cotyledonar leaf, in the form of a second and similar filiform leaf, to be followed by a third, and so on. The sheathing bases of these succeeding leaves become the coats of the Onion-bulb. The mternodes remain undeveloped until the plant is ready to blossom. Very similar is the germination of a date-seed, except that the protruding cotyledon does not lengthen so much, nor does it elevate the heavy seed. Instead of the seed being carried up, the lower end of the embryo, contain- ing the plumule, is pushed down more or less into the loose soil, from which in time the developing plumule emerges. 41. The embryo of Grasses, especially of those which yield the cereal grains, is more complex, owing mainly to the great de- velopment of the plumule and the manner in which c- its rudimentary leaves successively enclose each other. That of Maize or \\\ \i \:--:w.rs • ;i /^i 60 01 The floury part of the seed, which makes most of its bulk, is the albumen, largely composed of starch. The embryo is exterior to this, applied to one of its flat sides, and reaching from the thinner edge to or above the middle in the common variety of corn here represented. The form of the embryo is best shown, detached entire, in Fig. 58 : its structure appears in the sections. The outer part is the cotyledon, which incompletely enwraps the plumule : it adheres closely to the albumen by the whole back, and remains un- changed in germination : its function is to absorb nutritive Indian Corn, one of the largest, is most convenient for study. (Pig. 56-59.) FIG. 56. Section, flatwise, of a grain of Indian Corn, dividing the albumen and the embryo. 57. Similar section at right angles to the first. 58. A detached embryo : corresponding parts of Fig. 57 and 58 indicated by dottc-d lines. FIG. 59. Vertical section of Indian Corn across the thickness of the grain, dividing the embryo through the centre and displaying its parts : c, cotyledon ; p. plumule ; r, the radicle or caulicle. FIG. 60. Similar section of grain of rice. 61. Same of an oat-grain ; the parts as in Fig. 59. 26 MOUPHOLOiiY matter furnished by the albumen, and to transmit it to the growing plumule. The plumule consists of a succession of rudimentary leavo. sheathing and enclosing one another, on the summit of a very short axis, which is mainly the caulicle. otherwise called rad- icle. This is completely en- closed by a basal portion of the cotyledon and of the outermost leaf of the plu- mule, which form a peculiar sheath for it, named the Coleorhiza,1 i. c. root-sheath : consequently the first root or roots have to break through this covering. As in the Oak and Pea (32), the very first or outermost leaves of the plu- mule develop imperfectly and not into efficient foliage. The one in Fig. C2. which encloses the rest in the early growth, is left behind as a mere sheath to the base of the following and more perfect leaves : it is the same as the lowest in Fig. 63. The leaves are first developed : the internodes lengthen later, and the lowest lengthen very little. Not rarely the first root starts singly from the tip of the caulicle (Fig. 62, just as in Fig. 55) ; but others of equal strength follow from any part of the caulicle. and soon from the nodes above; and no tap-root is ever formed. 42. A Pseiido-monocotyfadonoits embryo occasionally occurs ; that is, one of the dicotyledonous type, of which one cotyledon is wanting through abortion. This occurs in Abronia. a genus related to Mirabilis. and bearing an embryo very similar to that represented in Fig. 17. IN. except that one cotyledon is absent. The anomaly of an acotyledonous embryo occurs in Dodder, a plant of the dicotyledonous type, but with both cotyledons 1 This, tlic t'li/inrliizi of Mirbcl. should not be confounded (as hy some it has been) with the "root-rap," or tissue which ordinary roots (whether primary or secondary) break through in their development or carry on their apex. Flci. •;.'. Early germination of Indian Corn. 63. More advaneeil germination of name: i-.inis prniliM -I'll tViim portion of stem above the cotyledon as well as below. OF THE ORGANS OF VEGETATION. 27 actually wanting, — a correlation with its parasitic mode of life. (64, Fig. 78.) 43. The dicotyledonous and the monocotyledonous character of the embryo is correlated with profound differences in the whole ulterior development, as revealed in the structure of the stem, leaves, and flower ; which differences mark the two great divisions of Phsenogamous plants, viz. DICOTYLEDONES or DICOTYLEDONOUS PLANTS, and MONOCOTYLEDONES or MONOCOTYLEDONOUS PLANTS, - names introduced into classification by Ray, and adopted by A. L. Jussieu, in his Genera Plantarum. CHAPTER III. MOEPHOLOGY AND STRUCTURE OF THE ORGANS OF THE PLANT IN VEGETATION. SECTION I. OF THE ROOT. 44. The Root, which has been called the descending axis, is that portion of the body of the plant which grows downward, ordinarily fixing the vegetable to the soil, and absorbing from it materials which the plant ma}r elaborate into nourishment. As alread}' stated (24), the root grows in length by continuous additions of new fabric to its lower extremity, elongating from that part only or chiefly ; so that the tip of a growing root always consists of the most newly formed and active tissue. It normally begins, in germination, at the root-end of the caulicle, or so called radicle. But roots soon proceed, or rna}' proceed, from other parts of the stem, when this is favorably situated for their production. The root does not grow from its naked apex, but from a stratum immediately behind it : consequently its blunt or obtusely conical advancing tip consists of older, firmer, and in part effete tissue. The tip of all secondary roots and rootlets FIG. 64. Magnified tip of root of a seedling Maple (such as in Fig. 9), sufficiently enlarged to indicate the cellular structure: a. the portion where growth is taking place; 6. the older and firmer tip. MORPHOLOGY <>!•' -1111. l:oOT. is similarly capped or protected.1 But the so-called rm>t-<-tiji is seldom so distinct or separable as to deserve a particular name. I.",. Nature of (irowlli. Cells. The development and growth of the root, as of other organs, results from the development, growth, and increase in number of certain minute parts, of which the plant is built up. These component parts are so much alike, at least in an early stage, and are so obviously formed all on one tvpe. that they take one common name, that of CELLS. These are the histological elements of plants. /. i: the units ,,| minute anatomical structure. While, in the morphology of the plant's obvious organs, analysis brings us to the j>/t>//n,i>er (Hi) as the individual element which by a kind of propagation produces its like in a second phytomer. remaining however in connection with the first, thus building up the general structure. so. in an analogous way, each of the obvious parts — each stalk or blade or rootlet — is microscopically determined to be com- posed of these ultimate organic units, generally called cells. The cell (cellulo. by the French conveniently termed cellule) is the living vegetable unit, in the same sense that the brick is the unit of a brick edilice. To make this analogy fairly complete, the bricks should be imagined to have a firm exterior or shell, and a soft or at length hollow interior, also to be living when incorporated into the structure, and finally to be produced in tin- forming structure by a kind of propagation. The production or increase in number of these cells by development from previous ones, and their successive increase in size up to maturity, are what constitutes vegetable ymx-ih.- The inspection through a 1 The notion that the tip of the root consists of delicate forming or newlv formed tissue, or bears some origan or structure of this nature la "Spongiol "). lias hardly yet been eliminated from the text-books and popular writings. It had no proper foundation in fact. In /.< mini, and in >ome other aquatics, and also in some aerial roots, this older lis-ue often separates into a real root-cap, free at base, like an inverted calyptra. - This. a< to the structure, is the subject of l/isln!,i-/i/ ; as to processes OF actions, the subject of I'/ii/aiiilnii/ .- both to be treated in a separate volume. I'll'.. ('..">. I'll'.. I'lM-li.iii- nl surf;!!1'1 'if Fi.it. i;t. more inaitriilii'il. rl.'.-irly cli*i>]:ivinit tin- >il:ir st run urr :ni. CM 1 Iril roof- hairs, wlii.'h ali'iiiinl on tin- H]I|MT part of Fiit <'.l. MORPHOLOGY OF THE ROOT. 29 simple microscope of a slender young root, and of thin slices of it immersed iji water, may serve to give a general though crude idea of the vegetable cellular structure, sufficient for the present purpose. Roots are naked; that is, they bear no other organs. When they send oil' branches, these originate from the main root just as roots originate from the stem ; and in both cases without much predetermined order. The ultimate and very slender branches arc sometimes called root-fibrils ; but these are only delicate ramifications of the root. Like any other part of the plant, however, roots may produce hairs or such like growths from the- surface, which are wholly distinct from branches. (383.) 41). Root-hairs. Roots absorb water, &c., from the soil by imbibition through the surface ; that is, through the walls of the cells, which are in a certain sense permeable to fluids, more readily when young and tender, less so when older and firmer. Roots, therefore, absorb most by their fresh tips and adjacent parts ; and these are continually renewed in growth and extended fur- ther into the soil. As the active surface of a plant above ground is enormously increased by the spread of foliage, so in a less degree is the absorbing surface of 3Toung roots increased by the production of root-hairs. (Fig. 64, upper part, and more magni- fied in Fig. 65, 6G.) These are attenuated outgrowths of some part of the superficial cells into capillary tubes (only one from each cell) , closed at the tip, but the calibre at base continuous with the cavity of the cell ; into which, therefore, whatever is imbibed through the thin wall may freely pass. These appear (as Fig. 64 shows) at a certain distance behind the root-tip. Further back the older or effete root-hairs die away as the cells which bear them thicken into a firmer epidermis. 47. To the general statement that roots give birth to no other organs, there is this abnormal, but 'by no means unusual excep- tion, that of producing buds, and therefore of sending up leafy branches. Although not naturally furnished with buds in the manner of the stem, yet many roots have the power of originat- ing them under certain circumstances, and some produce them habitually. Thus Apple-trees and Poplars send up shoots from the ground, especially when the superficial roots are wounded. And the roots of Madura or Osage Orange so readily originate buds that the tree is commonly propagated Ivy root-cuttings. 48. Kinds of Roots. The root, commonly single, which origi- nates from the embryo itself, is called the PRIMARY ROOT. (37.) Roots which originate from other and later parts of the stem, or elsewhere, are distinguished as SECONDARY ROOTS. But the 30 .MORPHOLOGY OF THE ROOT. hitter arc as normal as the primary root; that is, to stems so situated that they can produce them. Most creeping plants emit them freely, usually from the nodes : and so do most branches, not too old, when bent to the ground and covered with earth, thus securing the requisite moisture and darkness. Separate pieces of young stems (cuttings) can commonly be made to strike root. Upon this faculty of stems to originate roots depends all propagation by division, by laying or layering, by cuttings, ifcc. It is mainly annuals and common trees that naturally depend on the primary root; and most of these can In- made to produce secondary roots. Even leaves and leaf-stalks of some plants may be made to strike root and be used as cuttings. (77.) 49. Duration. By differences in respect to this, either the root or the plant, as the case may be. is distinguished into Annual, Biennial, or Perennial, according to whether life is contin- ued for a single year or season, for two, or for a greater number. The difference is not in all cases absolute or even well marked. 50. Annuals are plants which, springing from the seed, flower and seed the same year or season, and die at or before its close. They prod uce /irows roots, either directly from the embryo and succeeding joints of stem (as in Grasses, Fig. 63), or from a I n-rsistent primary or tap-root, more or less thickened into a trunk or divided into branches. The products of vegetation in all such herbs are not stored in subterranean or other reservoirs, but are expended directly in new vegetative growth, in the production of blossom, and finally in the maturation of fruit and seed. This completed, the exhausted and not at all replenished indi- vidual perishes. />!. But some annuals may have their existence prolonged by not allowing them to blossom or seed. Others, with prostrate stem or branches, may from these produce secondary roots, which, forming new connections with the soil, enable the newer growth to survive when the older parts \\ith the original root liave perished. And many herbs, naturally annuals, are continued from year to year through such propagation from the branches, used as layers or cuttings. Moreover, certain plants (such as Ricinus or Castor-oil Plant) , which are perennial or even arbo- rescent in warm climates to which they In-long. In-come annuals in temperate climates, early perishing by autumnal cold. .r>2. The annuals of cool climates, where growth completely ceases in winter, germinate in spring, mature, and die in or before autumn. But, in climates with comparatively warm and rain}' winter and rainless summer, many germinate in autumn, vegetate MORPHOLOGY OF THE BOOT. 31 through the winter, flower and seed in spring, and perish in early summer. These ma}- be termed WINTER ANNUALS. 53. Biennials are plants which, springing from the seed and vegetating in one season, live through the interruption of winter, and blossom, fructify, and perish in the next growing season ; their life being thus divided into two stages, the first of vegeta- tion, the second of fructification. In typical biennials, nearly the whole work of vegetation is accomplished in the first stage, with the result of accumulation of a stock of nutritive matter, to be expended in the second stage in the production of blossom and seed. This accumulation is usually stored in the root or in the base of a very short stem in connection with the root. The root of a biennial accordingly enlarges and becomes fleshy, or obese, as this matter accumulates. At the close of the growing season, — the leaves perishing and the stem having remained very short (with undeveloped internodes) , — the root, crowned with the bud or buds, contains the main result of the summer's work, AS provision for the next year's devel- opment and the completion of the cycle. This development, being thus amply provided for, is undertaken in spring with great vigor ; blossom, fruit, and seed are rapidly produced ; and the stock being consumed, but not at all replenished, the cells of the great root are now empty and effete, and the individual perishes. The Beet, Turnip, Parsnip, and Carrot are fa- miliar examples of biennials, with the store of nourishment in the root.1 The Kohl-rabi is a biennial with this deposit in the stem : the Cabbage, partly in the stem, partly in the head of leaves. 1 In these the caulicle enlarges with the root, so that the upper anil bud-bearing end is stem. Tap-roots of this kind are said, in descriptive botany, to be Fusiform or Spindle-shaped, when broader in the middle and tapering towards both ends, as in the common Radish (Fig. 67); Conical, when tapering regularly from base to tip, as in carrots, &c. ; Napiform, i. e. Turnip-shaped, when the thickened part is wider than high, &c. Fascicled Roots are those which form in clusters ; these may be slender or thickened. When much thickened, either irregularly or not of the above shapes, they are said to be tuberous. FIG. 67. Radish: a fusiform tap-root MORPHOLOGY <>F TIIK l;o<>r. ">l. But some plants, such as the Radish, which when they spring from seed in autuinii arc true liieiinials, will \vhcn raised in spring pass on directly to tlic (lowering stage in summer, or \vlicn sown after the warm season begins will often run through their eour>e as annuals. Then there are various liieiinials which thicken the root verv little and hold their leaves through the « C7 winter. Between these and winter annuals no clear demarcation can be drawn. As respects annual and biennial duration, the terms may for the most part be applied indiscriminately to the plant or to the root. We may say either that the plant is a biennial, or that its root is biennial. ")."). Perennials are plants which live and blossom or fructify year after year. They may or they may not have perennial roots. In trees and shrubs, also in herbs with growth from year to year from a strong tap-root, the root is naturally perennial. But in most perennials with only iibrous roots, these are produced anew from time to time or from year to }'ear. Also, \\hile some such roots remain fibrous and serve only for absorption, others may thicken in the manner of the ordinary biennial root and serve a similar use, /. e. become reservoirs of elaborated nourishment. The Dahlia (Fig. (IS) mid the Peony afford good examples of this. Sweet potato is another instance.1 Most such roots have only a biennial duration : the}- are produced in one growing season: they yield their store to form or aid the growth of the next. When perennials store up nutritive matter underground, the deposit is more commonly made in a subterranean portion of the stem, in tubers, conns, bulbs. &<•. (See 11 :»-! •_>:>.) ."><;. The distinction between annuals and biennials is at times so dillieult. and the particular in which they agree SO manifest. -namely, that of blossoming only once, then dying, as it were by exhaustion. — that it was proposed by I )e( 'andolle to unite 1 II is only by the readiness of this root to produee adventitious buds, especially from its upper part, that it lias been mistaken for a tuber, sueli as the eonnnon potato. Fit;. i;s. Kasrided and tuln-nnis i>r fusiform (secondary) roots of Dahlia: a, a. buds on liasr "I' thi' strm .MORPHOLOGY OF THE ROOT. 33 the two under the common appellation of MONOCARPIC plants, Plantce monocarpicce, taken in the sense of only once-fruiting plants ; and to designate perennials by the corresponding term of POLYCARPIC, Plantce poli/mr/iica;, literally many-fruited, taken in the sense of many-times fruiting.1 57. But the distinction even here is no more absolute than that between annuals and biennials. For example, it is not quite clear whether the Cardinal Flower and related species of Lobelia should be ranked as annuals, biennials, or perennials. The plants may blossom and seed toward the end of the season in which the}r came from seed : or, germinated in autumn, the small seedlings may survive the winter ; but whenever fructified the fibrous-rooted mother plant dies throughout ; yet usually not before it has established, and perhaps detached from the base, small offsets to blossom the next season; and so on. Then Honseleeks (Sempervivum) and such-like fibrous-rooted succu- lent plants multiply freely by offsets which are truly perennial in the sense that they live and grow for a few or several }*ears ; but when at length a flowering stem is sent up producing blos- som and seed, that plant dies as completely and in the same manner as any biennial, only the generation of offsets surviving. The same is true of the Century plant (Agave Americana, wrongly denominated American Aloe), which vegetates in the manner of the accumulating stage of a biennial, except that this continues for several or very many years, while the flower- ing stage, when it arrives, is precipitated and terminated in a single season. 58. Although the stem usually sends forth roots only when covered by or resting on the soil, which affords congenial dark- ness and moisture, yet these are in some cases produced in the open air. Roots may likewise subserve other and more special uses than the absorption of crude or the storing of elaborated nourishment. 59. Aerial Roots is a general name for those which are pro- duced in the open air. One class of these may serve the office of ordinary roots, by descending to the ground and becoming established in the soil. This occurs, on a small scale, in the stems of Indian Corn ; the lower nodes emitting roots which grow to the length of several inches before they reach the ground 1 These terms or some equivalents have a convenience in descriptive botany. But those employed by DeCandolle are not happily chosen, as has often been said. Mo»(florous (bearing progeny once) and Pohjtocous (bearing many times) would be more appropriate. 3 34 MOBPHOLOGY OF THE ROOT. into which they penetrate. More remarkable cases abound in those tropical regions where the sultry air. saturated with moist- ure for a large part ol'lhc year, favors the utmost luxuriance of vegetation. In the Palm-like Pandanus or Screw-Pine l (l-'i^. O V O ('•'.I), very strong root>, emitted in the open air from the trunk, and >ooii reaching the soil, give the appearance of a tree partially raised out of the ground. The fanioii> Iian\ an-tive of India (Fig. 71 ) is a still more striking illustration : for the aerial roots strike from the horizontal branches of tin- tree. often at a great height, at first swing- ing free in the air, but linally reach- ing and establishing themselves in the ground, where they increase in diameter and form accessory trunks, surrounding the original bole and supporting the wide-spread canopy of branches and foliage. Very similar is the economy of the Man- grove (Fig. 70), which forms impenetrable thickets on low and muddy sea-shores in the tropics throughout most parts of the world, extending even to the coast of Florida and Louisiana. Here aerial roots spring not only from the main trunk, as in tin- Pandanus, but also from the branchlets. as in the Hainan. Kven the radicle of the embryo starts into growth, protrudes, and attains considerable length while the fruit is still attached to the branch. .V.i'. Aerial Rootlets for (-limiting are familiar in the Ivy of tin- Old World (Iledera), Trumpet-Creeper (Teeoma radicans). and our Poison Ivy (Rhus Toxicodendron) : by the adhesion of 1 So named, not from :iny resemblance to a Pine-tree, but from a like- ness of the folia £i- to that of a Pine-Apple. FIG. 69. PaiKianus, or Screw-Pine; and in the background, 70, a Mangrove-tree (Khizopliora Manglr). MORPHOLOGY OF THE BOOT. 35 which the stems, as they grow, ascend walls and the trunks of trees with facility. Iii Rhus a superabundance of these rootlets is produced, thickly covering all sides of the stern. 60. Epiphytes or Air-Plants also have roots which arc through- out life unconnected with the ground. Epiphytes, or Epiphytic plants, as the name denotes, are such as grow upon other plants without taking nourishment from them. Deriving this from the air alone, they are called Air-plants. This name might be extended to the same or other kinds of plants attaching them- selves to bare walls, rocks, and the like, and unconnected with the soil, though such would not technically be epiphytes. Very many Lichens, Mosses, and other plants of the lower grade, and not a few phsenogamous plants, are in this case. The greater part of the phaenogamous Epiphytes pertain to two mouocotyle- donous orders, the Orchis family and that to which the Pine- Apple belongs, viz. the Bromeliacese. Their thread-like or cord-like simple roots either adhere to the bark of the supporting tree, securing the plant in its position, or some hang loose in the air. Of these, Orchids, i. e. plants of the Orchis family, are the most show}' and numerous, and of the greatest variety of forms, especially of their blossoms, which are often bizarre and fantas- tic. The}' belong, naturally, to climates which are both warm and humid ; the}' are highly prized in hot-house cultivation ; and, along with the hardy and terrestrial portion of the order, they are peculiarly interesting to the botanist on account of the singular and exquisite adaptation of their flowers in relation to insects which A-isit them. In some the blossoms curiously FIG. 71. The Banyan-tree, or Indian Fig (Ficus Imlica). ob' MORPHOLOGY OF THK I:OOT. resemble butterflies or other insects; as, for example, Oneidium 1'apilio. Fig. 1'2. Epiphytic orchids arc indigenous to the United States only from < ieorgia to Texas, and only in humble forms, in company with species of Tillandsia, representing liromeliace- ous epiphytes. The commonest of the latter tribe, and of most northern range, is the T. usneoides, the so-called Long Moss, which, pendent in long and tangled gra}" clusters or festoons from the branches of the Live-Oak or Long-leaved Pine, gives such a peculiar and sombre aspect to the forests of the warmer portions of our Southern States. 61. Parasitic Plants have the peculiarity that their roots, or what answer to roots, not only lix themselves to other plants, luit draw therefrom their nourishment, at least in part. Among crvptoii'anious plants very many Fungi art' parasitic upon or within living plants or animals. But only ph;cnogamous para- sites arc here under consideration. These may be divided into two classes: those with and those without green foliage. 62. (.iTni Parasites may be cither wholly or partially parasitic ; that is. they may draw all their support from a foster plant, or i'ic,. 7i'. Oncidium Papillo, and, 73, Comparettla roeea ; two showy epiphytes of the Orchis family; showing tin- moilc in which I lirsu Air-plants grow. MORPHOLOGY OF THE ROOT. 37 they may be likewise rooted in the soil, and receive from it materials of their food. Having green foliage, the}- are capable of elaborating such food, whether taken directly from the soil or from the crude sap of the foster plant. The Mistletoes (Viscum and its allies) are the principal examples of complete green parasitic plants. Seeds dropped b}^ birds on the boughs of trees germinate there ; the root-end of the caulicle points thither instead of towards the earth ; the root, or what would be such, pene- trates the bark and in- corporates itself with the sap-wood so perfectly that the junction of par- asite with foster trunk is like that of branch with parent trunk. The parasite is probably fed by both elaborated and crude sap, that is, both by what the foster tree has assimilated and what it has merely taken from the soil and air : the former it can at once incorporate ; the latter it has first to assimilate in its own green leaves. Sometimes one Mistleto is parasitic upon an- other of the same or of a different species. 63. Partially parasitic plants (mostly green) may be either woody and arborescent or herbaceous. The species of Clusia in tropical FIG. 74. Native epiphytes of Georgia, &c. : the erect one at the right an Orchid, Epidendrurn conopseum ; the hanging one Tillandsia usneoides, called Long Moss. FIG. 75- Boots of Gerardia flava : some of the rootlets attaching themselves para- sitically to the root of a Blueberry. (From a drawing by Mr. J. Staufier .) MORPHOLOGY OF THE HOOT. America (called Cursed Fig) are examples of the former. They form trees, send down aerial roots in tin- manner of the Banyan ; but, while some roots seek the ground, sonic may attach them- schcs to other trees parasitic-ally, and draw from them a portion of their support. The parasitism of certain herbaceous plants •with green foliage is clandestine, the connection being under- ground and therefore long unsuspected. This occurs in species of ( ierardia (at least of the section 1 )a.-\ sioma ) and other plants of the same family, the unciiltix ability of which is thereby explained. Also in Comandra and in their relatives the Thesinms of the Old World, belonging to a natural order (the- Santalacea-) which has much allinily with the entirely parasitic order ( Loranthaceie ) to which the Mist let o bi'lungs. 64. Pale or Colored Parasites, such as Beech-Drops, Pine-Sap. &c., are those which are destitute of green herbage, and are usually of a white, tawny, or reddish hue-; in fact, of any color except green. These strike their roots, or sucker-shaped di^-s. into the bark, mostly that of the root, of other plants, and thence draw their food from the sap already elaborated. They have accordingly no occasion for digestive organs of their own, i. e. for green foliage. The Dodder (Fig. 77) is a common plant of this kind which is parasitic above ground. Its seeds germinate in the earth, but form no proper root: when the slender twining stem reaches the surrounding herbage, it forms suckers, which attach surface of the supporting plant. 77 themselves 78 79 firmly to tin- penetrate its epidermis, and feed upon its juices; while the original root and base of the' stem perish, and the plant has no longer any connection with the soil. Tims stealing its nour- FIG. 76. Section of one of tin- attached rootlets of Geranlia. showing the union. FICi. 77. The common Dodder of thr Northern States (Cnscuta ciroiiovii), of the natural size, parasitic upon the stem of an herb: the uncoiled portion at the lower end shows the mode of its attachment. 78. The coiled embryo taken from the seed, con- sisting of naked caulicle and plumule; moderately magnified. 70. The same in germi- nation, elongating into a thread-like Icallrss stem. MORPHOLOGY OF THE EOOT. 39 ishment ready prepared, it. requires no proper digestive organs of its own, and, consequently, does not produce leaves. This economy is foreshadowed in the embryo of the Dodder, which is a naked thread spirally coiled in the seed (Fig. 78, 79), and presenting no vestige of cotyledons or seed-leaves. A species of Dodder infests and greatly injures flax in Europe, and some- times makes its appearance in our own flax-fields, having been introduced with the imported seed. Such parasites do not live upon all plants indiscriminately, but only upon those whose elaborate juices furnish a propitious nourishment.1 Some of them are restricted, or nearly so, to a particular species ; others show little preference, or are found indifferently upon several species of different families. Their seeds, in some cases, it is said, will germinate only when in contact with the stem or root of the species upon which the}* are destined to live. Having no need of herbage, such plants may be reduced to a stalk bearing a single flower or a cluster of flowers, or even to a single blossom developed from a bud directly parasitic on the bark of the foster plant. Of this kind are the several species of Pilostyles (para- sitic flowers on the shoots of Leguminous plants) in Tropical America, one species of which was discovered by Dr. Thurber near the southern borders of New Mexico. Its flowers are small, only about a quarter of an inch in diameter. The most wonderful plant of this kind is that vegetable Titan, the Raf- flesia Arnoldi of Sumatra (Fig. 80) which grows upon the stein of a kind of a Cissus or Vitis. It is a parasitic flower, measuring nine feet in circumference, and weighing fifteen pounds ! Its color is light orange, mottled with yellowish- white. 1 Monotropa or Indian Pipe (and perhaps some related plants), although probably parasitic on living roots in early growth, appears to live afterwards in the manner of the larger Fungi, upon leaf-mould and decaying herbage. Its mode of life should be investigated. FIG. 80. Rafflesia Arnoldi ; an expanded flower, and a bud, directly parasitic on the stem of a vine: reduced to the scale of half an inch to a foot. 40 MOKPHOLOGY OF BUDS. SECTION II. OF BUDS. 65. Buds are the genus of stems : they are axes with their appendages in an e:irly state. LEAF-BUDS (tii.MM.K) are those devoted to vegetation, and tin- parts, or some of them, develop as leaves. MI.\KI> m i>s contain both foliage and (lower or flowers. Fi.owi.K-r.i i» ( Ai. AI;A>TKA) are iniexpanded blossoms. These arc considered in another chapter. 66. The conspicuous portion of an ordinary bud, or that which first develops, usually consists of leaves, or scales the hoinologues of leaves ; the axis itself being very short and undeveloped. If this remains comparatively short, the leaves as developed are crowded in a rosette, as in a Ilouseleek (Fig. (J1'(). a Barberry and the Larch: when the internodes lengthen, the lea\es are interspaced upon the axis. G7. The cotyledons and plumule of the embryo are. morpho- logically, the first bud, on the summit of the initial stem, the caiilicle. This in germination and subsequent growth develops into a leafy stem, in the manner already described. Normally this stem has the capacity of growing on in this way from the apex or growing point, which is always potentially a bud, the njiicalov terminal bud (15). Sometimes it is merely potential. and there is no external structure visible until the new growth begins, or the bud is said to be latent. 68. But commonly, in plants that live from year to year, growth is divided into seasons or stages, with intervals of repose. In such cases, especially in trees and shrubs, instead of a continuous succession of foliage, the period of interruption is apt to be ma iked by the production of scales (End-scales, Perulce, etc.) or dry teguments, which serve to protect the tender rudiments or growing point within during the season of rest. This being the winter-season in cold climates, Linn;i?us gave to such bud-cover- ings the common name of HlBERNACULUM. From the usually st/innnose (scale-like) character of this covering, such buds lake the name of 69. Scaly Buds. Large and strong ones of this kind, such as those of Ilorsechesl nut. Magnolia. Hickory. Lilac. &c., may In- taken as the type of bud. The scales serve to protect the ten- der parts within against injury from moisture and from sudden changes in temperature during the dormant or earliest growing state. To ward off moisture more etlectually. they are sometimes coated with a waxy, resinous, or balsamic- exudation, as is con-1 spicuous on the scales of the Ilorsechestnut, Balsam-Poplar or Balm of Gilead. and Balsam-Fir. To guard against sudden MORPHOLOGY OF BUDS. 41 changes of temperature, they are often lined, or the rudimentary leaves within invested with non-conducting down or wool. 70. Nature of Bud-scales. That they answer to leaves is made manifest by a consideration of their »i situation and arrangement, which* are the same as of the proper leaves of the species ; and by the gradual transi- tions from the former to the latter in man}' plants. In the Turions, or sub- terranean budding shoots of numerous perennial herbs, and in the unfolding buds of the Lilac and Sweet Buckeye (JEsculus parviflora), every gradation may be traced between bud-scales and foliage, showing that no line of distinc- tion can be drawn between them, but that the two are essentially of the same nature, are different modifications of the same organ. In the Lilac they may be regarded as the blade of the leaf, modified and depauperate ; in the Buckeye (Fig. 233), and therefore in Horsechestnut, as the base of leaf-stalks ; in Magnolia (Fig. 81, 82), in the Tulip-tree, and in the Beech, they are evidently stipules. They must therefore be referred to in the section on the morphology of leaves. (227.) 71. Naked Buds, &c., of shrubs and trees, even in climates with severe winter, are not unknown, that is, buds unprotected by special scales or other coverings. For example, the latest pair of leaves of the season in Viburnum nudum, V. lantanoides (Hobblebush) , and the like, remain in a nascent state over winter without covering, and ex- pand into the first foliage in the spring. Yet V. Opulus (Snowball, &c.), another species of the same genus and inhabiting the same region, has well-formed scaly FIG. 81. Branch of Magnolia Umbrella, of the natural size, crowned with thu terminal bud; and below exhibiting the large rounded leaf-scars, as well as the rings or annular scars left by the fall of the bud-scales of the previous season. 82. A detached scale from a similar bud ; its thickened axis is the base of a leaf-stalk ; the membranous sides consist of the pair of stipules united with it. 42 MORPHOLOGY OF BUDS. b ~ leal-buds. In other hardy shrubs and trees, the buds, equally or almost destitute of scales, are minute, hidden in or un- der the bark, or otherwise inconspicuous until vernal growth commence-. Phila- delphus and Taxodimn are of this kind. 7~2. Subpetiolar IJuds. Smiie leaf-buds are singularly cov- ered in their early state and through the summer, as iu the Locust (Robinia). Honey-Lo- cust Fig. !MJ (where they re- main very undeveloped), in Yellow AYood (C'ladrastis).and more conspicuously in the Plane-tree i Platauus. Fig. -S7 i : here they are all formed tin- der the base of the protecting leaf-stalk, which in Plane-tree forms a sheath or inverted cup. as st ss very like a candle-extinguisher. fitted to and concealing the conical bud until autumn, when hy the fall of the leaves these buds are exposed. FIG. s:: Diagram <>f vertical section nf a strong lunl. such as of Horseclifstiiut. l. 'I'ln- :i\is iif tin- S:IIIK> ilrvcln]ii!i;:, tin- flniiLMt i"ii lif^iiiiiint: «'itli tlir l"\vr>t intcr- r, sunn I'n lli < \vc Fin'. *1. *•">. '-'I ). this is commonly the strongest, or among the Stronger. But in many eases it habitually or eommonly fails to appear. In the Elm (with leaves and therefore bud> alternate). the bud axillary to the last leaf of the season takes its place. In the common Lilac, a pair of buds, which were in the axils of the uppermost of the (opposite) leaves, seem to replace the terminal bud, which seldom develops. (Fig. «G.) When all the regular buds make their appearance, and the leaves are opposite, the stem \\ill be crowned with the terminal bud, having an axil- lary bud on each side of it. (Fig. 88.) 7('.. Accessory IJuds. These are. as it were, multiplications of the regular axillary bud. giving rise to two, three, or more, instead of one ; in some cases situated one above an* other (superposed), in others placed side by side (collateral). In the latter case, which occurs occasionally in the Hawthorn, in cer- tain Willows, in the Maples (Fig. 88), &c.. the axillary bud seems to divide into three. or itself to give rise to a lateral bud on each side. On some shoots of the Tartarean Honeysuckle (Fig. 90) from three to six buds appear in each axil, one above another, tin- lower being successively the stronger and earlier produced ; and the one- immediately in the axil, therefore, grows in preference : occasionally two or more of them grow, and superposed accessory branches result. It is much the same in Aristoloehia Sipho, except that the uppermost bud is there1 strongest. i. 88. Branch of K«'<1 Maple, at the middle bearing triple axillary bads, placed siilc liy siili;. FIG. 89. Pice.' of ;i l.nmrli of the Butternut, with arn-ssory l.u.ls pl.-uvd one above anoili.T: ft, tin- leaf si'iir: /', proper axillary Innl: c, n«ls. KM '• '.MI. Part of a l>r:ui< li of Tartarean Honeysuckle, with crowded accessory bud8 superposed in the axil of each leaf. MORPHOLOGY OF STEMS. 45 So it is in the Butternut (Fig. 89), where the true axillary bud is minute and usually remains latent, while the accessor}' ones are considerably remote, and the uppermost, which is much the strongest, is far out of the axil : this usually develops, and gives rise to an extra-axillary branch. 77. Adventitious Buds are such as are abnormal and irregular, being produced without order and from any part of the stem, or even from roots. The latter, like the mternodes of a stem, although normally destitute of buds, do produce them notwith- standing in certain cases, especially when wounded, and in some plants (such as Blackberries) so freely that gardeners propagate them by root-cuttings. The steins share this tendency; and buds are apt to break out on the sides of trunks, especially when wounded or pollarded, or to spring from new tissues produced on cut surfaces, especially where the bark and wood join. Even leaves may develop adventitious buds, and then be used for propagation. In Bryophyllum, such buds, followed by rootlets, are freely produced on the margins of the blade or of its leaflets. In Begonia, a leaf, used as a cutting, will root from the base of the petiole stuck in the soil, and produce buds on the blade, at the junction with the petiole, or elsewhere. SECTION III. OF THE STEM. § 1. GENERAL CHARACTERISTICS AND GROWTH. 78. The Stem is the ascending axis, or that portion of the trunk which in the embryo grows in an opposite direction from the root, seeking the light, and exposing itself as much as pos- sible to the air. All phsenogamous plants possess stems.1 In those which, in botanical descriptions, are said to be acaulescent, or stemless, it is either very short, or concealed beneath the ground. Although the stem always takes an ascending direction at the commencement of its growth, it does not uniformly retain it ; but sometimes trails along the surface of the ground, or burrows beneath it, sending up branches, flower-stalks, or leaves into the air. The common idea, that all the subterranean portion of a plant belongs to the root, is incorrect. Equally incorrect is the common expression that plants spring from the root. Roots spring from the stem, not the stem from the root. (21, 24, 37, 44.) 1 There are, however, reduced forms in which there is no distinction of axis and foliage ; but most of these are clearly leafless rather than stemless, and not even in Lemna and Wolffia can the stem be said to be wanting. 46 .MORPHOLOGY OF STEMS. 79. While the root normally gives birth to no other organs, but itself performs those functions which pertain to the relations of the vegetable with the soil. — binding it to the earth and absorbing nourishing materials from it. — the aerial functions of \i--etation are chielly carried on. not so much by the stem it- self as by a di-tind set of organs which it bears, namely, the leaves. Hence, the production of leaves is one of the charac- teristics of the stem. These are produced only at certain definite and symmetrically arranged points, called nodes. (!:'.. •_':'..) 80. Development and Structure. In a bud or undeveloped stem. the nodes are in contact or close proximity. In the develop- ment, growth in length takes place in such manner as to carry these apart more or less, according to the degree of elongation, that is. the internodes (13) elongate. The order of development is from below upward, the lowest internode first lengthening, the others in regular succession. Each completes its growth. with more or less rapidity, although the length attained varies greatly in different stems, in different parts of the same stem, ami under different conditions. I'lilike the root, in which the elongation of formed parts is very soon finished and therefore only the tip is perceptibly growing, internodes go on growing throughout, and several formed internodes may be growing simultaneously, thus producing elongation throughout a consid- erable extent of stem and with considerable rapidity. But each internode grows independently. Some parts of an internode mav lengthen faster or continue in growth longer than others; this is usually the upper portion, at least in long internodes and when every part is equally exposed to light. 81. The development of a stem from a bud is wholly like that from the embryo, and has already been described in Chap. II. It exhibits similar variations as to rapidity and vigor, dependent upon the constitution of the bud, — which, like the plumule in the seed or seedling, may be either latent or much developed lie fb re growth begins. - -also upon the amount of nourishment provided. Strong buds commonly have their parts, or some of I hem. ready formed in miniature, and a store of elaborated nour- Miment in the parent stem to draw upon. Those well-developed buds which in many of our shrubs and trees crown the apex or occupy the axils of stem and branches early in the preceding summer (as in .Magnolia. Fig. *1. llorsechestnut, Fig. *•">. and Hickory, Fig. '.)!) often exhibit the whole plan and amount of the next year's growth; the nodes, the leaves they bear, and sometimes the blossoms being already formed, and only requiring the elongation of the internodes for their full expansion. As CHARACTERISTICS AND GROAVTH. 47 the bud is well supplied with nourishment in spring by the stem on which it re'sts, its axis elongates rapidly ; and although the growth commences with the lowest internode, yet the second, third, and fourth internodes may begin to lengthen long before the first has attained its full growth. Such very strong buds are usually terminal ; but sometimes, as in Lilac (Fig. 86), they are the uppermost axillary, which take the place of a suppressed or abortive terminal bud. 82. Such wood}' stems, developed from a strong bud, and terminated at the close of the season's growth by a similar bud, may be continued from year to year in an unbroken series. A set of narrow rings on the bark (Fig. 85 a] commonly marks the limit of each year's growth. These are the scars left by the fall of the scales of the bud ; and these, in the Horsechestnut, and in other trees with large scaly buds, ma}' be traced back on the stem for a series of years, growing fainter with age, until the}' are at length obliterated by the action of ^ the weather and the distention caused by the increase of the stem in diameter. The same is the case with the more conspicuous Leaf-scars, or marks on the bark left by the separation of the leaf-stalk, which are for a long time conspicuous on the shoots of the Horsechestnut (Fig. 85 &), the Magnolia (Fig. 81), and Hickory, Fig. 91. 83. Ramificatiou. BRANCHES (14-16) are secondary stems developed from a primary one, or tertiary ones from these, and so on. Ultimate or small ramifications of latest order are some- times called BRANCHLETS. The terminal bud continues the stem or axis which bears it. Lateral buds give rise to branches.1 As the normal lateral buds are axillary (75), so are normal branches. The symmetry or arrangement of branches, being that of the buds from which they are developed, is fixed by and follows that of the leaves. When the leaves are alternate, the 1 Dichotomy or forking, the division of an apex into two, although of com- mon occurrence in the lower cryptogamous plants, occurs so rarely and exceptionally, if at all, in phasnogamous plants that it may here be left out of view. In phasnogamous plants only the ramification of axes should take the name of branches. That is, roots and stems branch ; and the term may without confusion be extended to hairs and all TRICHOMES (383) when com- pound, but not to leaves and their modifications. FIG. 91. End of a Hickory branch (Carya alba), with a strong terminal and smaller axillary buds. 48 MORPHOLOGY OF STEMS. branches will l>c alternate ; when the leaves are opposite, and the buds develop regularly, the: branches will be opposite. &c. This holds in I'aet siilliciently to detenniiie and exemplify the plan of ramification ; but, if entirely carried out, there would be as many branches as leaves. This could rarely if ever be, even in primary ramilieat ion. •si. Non-development of Buds. Some of the buds are latent or merely potential, that is. do not make their appearance : of those which do appear only a part actually grow into branches : and of these some are apt to perish at an early stage. In our trees, most of the lateral buds generally remain dormant lor the first season : they appear in the axils of the leaves early in summer, but do not grow into branches until the following spring; and even then only a part of them grow. Sometimes the failure occurs without appreciable order; but it often is nearly uniform in each species. Thus, when the leaves are opposite, there are usually three buds at the apex of a branch ; namely, the terminal, and one in the axil of each leaf; but it seldom happens that all three develop at the same time. Sometimes the terminal bud continues the branch, the two lateral generally remaining latent. as in the I lorsecliestnut (Fig. 85) ; sometimes the terminal one fails, and the lateral ones grow, when the stem annually becomes twro-forked, as in the Lilac, Fig. 86. The undeveloped buds do not necessarily perish, but arc ready to be called into action in case the others are checked. When the stronger buds are destroyed, some that would else remain dormant develop in their stead, incited by the abundance of nourishment, wliieh the for- mer would have monopoli/ed. In this manner our trees are soon reclothed with verdure, after their tender foliage and branches have been killed by a late vernal frost, or consumed by insects. And buds which have remained latent for several years occasion- ally shoot forth into branches from the sides of old stems, especially in certain trees. 85. .Most branches springing from old trunks, however, as in Willows and Poplars, especially when wounded or pollarded, originate from adventitious buds (77), which occur without order. So also when accessory buds (7<>) develop into branches, normal symmetry is more or less disturbed, as by contiguous shoots standing directly over each other in Tartarean Honey- suckle, or by a branch far out of the axil in Walnuts (Fig. 89) and Honey-Locust. Fig. !•(>. 86. Exeiirreiit and Deliquescent Steins. Sometimes the primary axis is prolonged without interruption, even through the whole life of a tree (unless accidentally destroyed), by the continued CHARACTERISTICS AND GROWTH. 49 evolution of a terminal bud, or by some upper strong bud which equally becomes a leader, — forming an undivided main trunk, *from which lateral branches proceed ; as in most Fir-trees. Such a trunk is said to be excurrent. In other cases, the main stem is arrested, sooner or later, either by flowering, by the failure of the terminal shoot, or by the more vigorous develop- ment of some of the lateral buds ; and thus the trunk is dissolved into branches, or is deliquescent, as in the White Elm and most of our deciduous-leaved trees. The first naturally gives rise to coni- cal or spire-shaped trees ; the second, to rounded or spreading forms. As stems extend upward and evolve new branches, those near the base, being overshadowed, are apt to perish, and thus the trunk becomes naked below. This strikingly occurs in the excurrent trunks of Firs and Pines, grown in forest, which seem to have been branchless to a great height. But the knots in the centre of the wood are the bases of branches, which have long since perished, and have been covered with a great number of annual layers of wood, forming the clear stuff of the trunk. 87. Definite and Indefinite Annual Growth of Branches. In many of our trees and shrubs, especially those with scaly buds, the whole year's growth (except on certain vigorous shoots) is either already laid down rudunentally in the bud, or else is early formed, and the development is completed long before the end of summer ; when the shoot is crowned with a vigorous terminal bud, as in the Horsechestnut (Fig. 85) and Magnolia (Fig. 81), or with the uppermost axillary buds, as in the Lilac (Fig. 86) and Elm. Such definite shoots do not die down at all the follow- ing winter, but grow on directly, the next spring, from these terminal or upper buds, which are generally more vigorous than those lower down. In other cases, on the contrary, the branches grow onward indefinitely, until arrested by the cold of autumn : the buds at or near their summit are consequently young and miniatured, or at least the lower and older axillary buds are more vigorous, and alone develop into branches the next spring ; the later-formed upper portion most commonly perishing from the apex downward for a certain length in the winter. The Rose and Raspberry, and among trees the Sumac and Honey- Locust, are good illustrations of this sort ; and so are most perennial herbs, their stems dying down to or beneath the sur- face of the ground, where the persistent base is charged with vigorous buds, well protected by the ground, for the next year's vegetation. 88. Man)' of the details and applications of ramification, of most importance in morphology and descriptive botany, relate .".'I MdliniOLOGY OF STK.MS. to :mtlii>t:i\y or inllorescence (Cli;i[). V.), which has its own terminology. But sonic «>!' its term- may be conveniently employed in the description of ramification unconnected with* flowering. '&• § 2. FOI:M- OF STEMS AND BKA.NCIIES. 89. On the sixe :md duration of the stem the oldest and most obvious division of plants is founded, namely, into JlerK-. Shrubs. ;md Trees. !HI. Herbs :uv plants in which the stem does not become woody and per-i-teiit , but dies annually or alter flowering, down to the ground at least. The dillereiice l>ct\veeii minimi. lii,-n n'ml . and i>cri'iin!(il herbs has already been pointed out in the chapter on the root (."»()-.") 7 ) . and the gradations between them indicated. Herbs pax into shrubs and shrubs into trees through every gra- dation. The following delinitions are thcrelbre only general: — !)1. I'lulershrubs, or Xi/Jfr/i/imsi- plants, are \voody plants of hunilile stature, their stems rising little above the surface. If less decidedly woody, they are termed Sllffrutescent. '.)•>. Shrubs are woody plants, with stems branched from or near the ground, and less than live times the height of a man. A shrub which approaches a tree in si/.e. or imitates it in aspect. is said to In- Arborescent. !>.'!. Trees are woody plants with single trunks, which attain at least four or five times the human stature. Yet the name of tree is not to be denied to a woody plant having a >ingle and stout trunk of less altitude ; and those which grow in a bushy manner, sending up a cluster of stems from the ground to the height of thirty feet or more, may -till lie called shrubs. '.i|. The erect position, elevation above the soil, and self-sup- port, are normal conditions of the stem, but are far from universal. And certain kinds of stem or branches are sulliciently peculiar to have received substantive names : other equally peculiar forms have no special names. There are, moreover, certain organs (such as spines and tendrils) which are commonly homologous (!•_') with stems, but not always. Two kinds of erect stems have special names in descriptive botany. It."). Culm is a name applied to the peculiar closed-jointed stem of Grasses and Sedges, whether herbaceous, as in most (irasses, or woody or arborescent, as in the Bamboo. 96. CauuYx is the name technically applied to the trunk of Palms (Fig. 1-M'>). Tree-Ferns, and the like, consisting of a commonly simple column, the surface beset with scales, — the SPECIAL FORMS. 51 bases of former leaf-stalks, — or marked by scars, left by their fall. This name was used b}' botanists anterior to Linnaeus for any tree-trunk, but is now used for the peculiar stems above- mentioned ; also for the persistent base of a stem, otherwise annual, which throws up fresh herbaceous stems or stalks from year to year. Such short and enduring stems, being usually near the ground or under it, were commonly mistaken for roots. The old English name of Stock is sometimes used in botanical description for all short and enduring sterns of this sort, whether rising somewhat above or concealed beneath the surface of the soil. 97. A Scape is a stem or branch which rises from beneath or near the surface of the ground and bears flowers, but no proper foliage. It therefore belongs to inflorescence. (265.) Scapes usual!}' spring from some one of the subterranean forms of stem. 98. Of stems which do not stand upright in the air there are various modifications and gradations. 99. Scandent or Climbing Steins are those which rise by attaching themselves to some extraneous support. This is effected in various ways ; in some by the action of the stem itself, in others by that of organs which it bears.1 100. Voluble or Twining Steins, or Twiners, are those which ascend by coiling round a support, which must accordingly be comparatively slender, or at least not too large. Some ascend by coiling twwith the sun" (that is, from right to left of the observer viewing the coil from the outside 2) , as the Hop ; more, 1 See Darwin, The Movements and Habits of Climbing Plants, London and New York, 1875. Also the earlier paper on the subject in Journal of the Linnean Society, ix. 1865. Note that in North America climbing plants in general are in popular language called Vines (e. g. Hop- Vine, Grape- Vine, Squash- Vine, &c.), a name which properly belongs to Vitis only. 2 Dextrorse and Sinistrorse, i. e. to the right or to the left, are almost indis- pensable terms, but there is an ambiguity and discrepancy in their use. Darwin (in Climbing Plants, above referred to) seeks to avoid this by usually employing the terms " with the sun," and " against the sun," phrases which would be unmanageable in terminology. The writer (in Amer. Jour. Sci. ser. 3, xiii. 391) suggested Eutropic for the former, Antitropic for the latter, to be used in case it is preferred to evade rather than to encounter the ambiguity. Probably the terms dextrorse and sinistrorse, or right and left, will continue in use, as most natural and convenient. Now, in the first place, it should be understood that a plant, or at least a plant's axis, having no front and back, can have no right and left of its own. These relations of direction must refer to the right and left of an observer. All depends, accordingly, upon the position which the viewing observer is supposed to occupy when he predicates the direction of the turns of a helix or of the over- lapping of the parts of a bud. Linnaeus supposed the observer to view the 52 MORPHOLOGY OF STEMS. 91° by coiling in the opposite din-diem, as the Bean (Phaseolus), the woody Aristolochia Sipho, the Morning Glory (Fig. 91°) any attachment, through the development of sucker-like discs, along the whole contiguous surface. (Fig. 77.) The various actions through which plants elimb, and the attendant phenomena, are physiological, and will be treated in the second part of this Text Book. The most complete and satisfac- tory discussion of the subject, of a readable sort, is that of Darwin's volume, referred to in a preceding note. 101. Leaf-Climbers are those in which support is gained by the action, not of the stem itself, but of the leaves it bears ; in most by the coiling or clasping of petioles, as in Clematis, Maurandia, Tnipa-ohini, and Solanum jasminoides (Fig. 235) ; in some by the incurvation of leaf-blades or portions of them, as in Adlu- mia : or by an extension of the midrib into a hook or short ten- dril, as in Gloriosa ; or by the transformation of some of the blades of a compound leaf into hooks or tendrils, as in C'oba-a and the Pea. 102. Tendril-Climbers (Fig. D2-95) are those in which the prehension is by a tendril, a slender filiform body, either simple or branched, specially adapted to the purpose, and capable of coiling, either to secure a hold, or to draw the stein up to the coil or circle from the inside: Molil, Palm, Braun. and the l)e( 'andolles adopt this, and the latter insist on it. Such authority should lie decisive, it" (•(HiiiiKiii usaue and popular sense \\ent along with it. But some of the botanists following Linna-us adopted the reverse view ; and to the promt writer, as to Bcntham and Hunker, Darwin, Kichler, and in part pxtrornely twining stem of Morning Glory, Ipomoea purpuroa. SPECIAL FORMS. 53 support. In certain tendrils the attachment to the support is by a sucker-like disc at the apex, as in the Virginia Creeper or Ampelopsis, Fig. 94. 103. Root-Climbers are those in which the stems produce aerial rootlets (59a), which fix themselves to a supporting surface along which the stem creeps or ascends. In this way Trumpet Creeper (Tecoma radicans), Ivy, and Poison Ivy (Rhus Toxi- codendron) climb extensively. 104. Steins or branches which neither climb nor stand upright may have their direction or habit of growth expressed by certain adjective terms ; such as Ascending or Ass-urgent, when they rise obliquely upward ; Reclining, when from an ascending or erect base the upper part recurves and trails ; Decumbent, when trailing along the ground, but with apex assurgent ; Procumbent or Prostrate, when lying at length upon the ground ; Repent or Creeping, when growing prostrate on the ground and rooting as they grow. Also applied to similar stems grow- ing under, as well as upon the surface of the soil, as in Couch- Grass and Mint, Fig. 99. 105. A Sucker (Surculus) is an ascending stem rising from a subterranean creeping base. The Rose and Raspberry multiply freely b}T suckers. Such plants are easiest to propagate " by division." 106. A Stolon is a prostrate or reclined branch which strikes root at the tip, and then develops an ascending growth, which becomes an independent plant. 107. An Offset is a short stolon or a short sucker. Houseleek (Fig. 9P) offers a familiar example. By offsets, some herbs, otherwise annuals, are continued from year to year in a vegetative progeny (Lobelia cardinalis, &c.), and peren-i nials may thus establish colo- nies around a parent individual. 108. A Runner (Flagellum) is a filiform or very slender stolon, naked and tendril-like except at tip, where it roots, develops a bud, and so a new plant. The Strawberry furnishes the most familiar example. FIG. 91*. Houseleek (Sempervivum tectorum) with offsets. 54 MORPHOLOGY OF STEMS. 109. The two following are organs which may be of axial nature, or may not. This may ordinarily be determined by posi- tion. Any direct continuation of stem or I »ranch must be of axial nature, that is. of the nature of stem ; and the same is true of whatever primarily develops in the axil of a leaf. Conversely, whatever subtends a lateral axis or branch may be taken for a leaf or foliar production, being in the place of such. 110. A Tendril, a thread-shaped and leafless body, capable of coiling spirally, and used for climbing (102) , is homologous with stem in Grape- vines (Fig. t>2) : for the uppermost tendril is seen to be a direct continu- ation of the stem. The small bud which appears in the axil of the uppermost leaf will in its growth produce another internode and leaf, or some species more than one, but will terminate in a similar tendril : the present terminal tendril will have then become lateral and opposite the leaf, like the three in the .lower part of I'K I. !ij. End of a shoot of the Grape-vine, with young tendrils: a sympodial (See note.) I'M* 1. 93. A portion of a stem of Ampelopsis quinquefolia, or Virginia Creeper, with a lent' :nid a ti-ndril. FIG. 94. Ends of the latter, enlarged, showing the expanded tips or discs by which they cling. SPECIAL FOKMS. 55 the figure.1 The tendrils of Virginia Creeper (Fig. 93) are of the same nature and position. But, instead of laying hold by a coiling of the tip, when it has reached an}' solid surface, such as a wall or tree-trunk, the tip expands into an adhesive disc, which forms a secure attachment. (Fig. 94.) In a related plant, Vitis (Cissus) tricuspidata of Japan, these disks terminate the branches of very short tendrils : consequent!}* the shoots as they grow are at once applied closely and secured firmly to the surface of the sup- port, — an admirable adaptation for climbing walls and trunks. 111. The simple tendril of a Passion- flower, being in the axil of a leaf (that is in the position of a branch) , is also of axial nature : it is a leafless and simple branch, composed of one long and slen- der internode, devoted to the purpose of climbing. Fig. 95 shows in all stages the admirably active tendrils of Passiflora sic}*oides. This is a Mexican species, remarkable for the rapidity and freedom with which the tendrils move. The lowest tendril in the figure is attached and coiled : the next is free and coiled in , one helix : the third is outstretched and seeking a support. For tendrils which are not homologous with stems, 95 see Sect. IV. 228. 112. A Spine or Thorn (Fig. 96, 97) is usually a branch or the termination of a stern or branch, indurated, leafless, and attenuated to a point. The nature of spines is manifest in the Hawthorn (Fig. 97), not only by their position in the axil of a leaf, but often by producing imperfect leaves and buds. And in the Sloe, Pear, &c., many of the stinted branches become spinose or spinescent at the apex, tapering off gradually into a rigid and leafless point, thus exhibiting every gradation between a spine and an ordinary branch. These spinose branches are less This forms what is called a St/mpodium or Si/?npodial stem, which is mor- phologically made up of a series of superposed branches. (Sec Chapter V. 281,282.) In contradistinction, a stem formed by the continued development of a terminal bud is Monopodial or a Monopodium. Fig. 95 is an example. FIG. 95. Leafy shoot of Passiflora sicyoides, of Mexico, with fixed and coiled, free and full grown, and forming tendrils. MORPHOLOGY OF STEMS. liable to appear on the cultivated tree, when duly cared for, such branches being thrown mostly into more vigorous growth. In the Hawthorn, the spines spring from the main axillary bud. while accessory buds (76), one on each side, ap- pear, and grow the next sea- son into ordinary branches. In the Honey-Locust, it is the uppermost of several ac- cessoiy buds, placed Car above the axil, that develops into the thorn (Fig. 00) . Here the spine itself usually branches, and sometimes becomes ex- tremely compound. 11:5. For spines which are homologous with leaves, or parts of a leaf, see Sect. IV. 227°. Prickles, such as those of Brambles and Roses, are superficial outgrowths from the bark, of a different nature (:>*.'!). and of small morpho- 97f' 114. Subterranean Stems are hardly less diverse than the aerial. The}- are classed as RHIZOMES, TUBEUS, COUMS, and BULBS, the forms passing one into another by gradations. 11;"). Rhi/oma (Hliizome, or in English ROOTSTOCK) is a gen- eral name for any horizonal or oblique perennial stem, which lies on the ground or is buried beneath its surface. It sends off roots of a fibrous or slender sort wherever it rests on or is cov- ered by llic soil, and usually produces from its apex some kind of aerial stem, cither lealy or as a flower-stalk (*>•:il. with the terminal bud, the base of tip' stalk of the season, and tlnvi- si-ars from whii-h the latter has separated in ai many former years. FIG. Idl. Khi/oma of Kiphylleia eymosa. showing six years' crowlh. ami a bud for tlir si-vcnlli: <(, Ihr luid: l>, base- of Ilir stalk of the curn-nt yrar: ,\ scar Irfi by tlm decay of the annual stalk of the year before; and beyond are the sears of previous years. KIG. IIIL-. Shoot ami yoiins; rootstock of Trillium erectum. with only terminal bud. SPECIAL FORMS. 59 species, and in older individuals, it is longer, often oblique, and branching, and bears the scars from which the annual aerial growths have separated.1 Nymphaea odorata, the sweet-scented white Water Lily, grows by very long, stout, and simple rootstocks. In N. tuberosa the sides of the roots tock produce short lateral branches or tubers. 117. A Tuber may be morphologically char- acterized as a short thickened rhizoma on a slender base, or a rootstock some portion of which — mostly a terminal portion and involv- ing several nodes — is thickened by the depo- sition of nourishing matter. A potato and a Jerusalem artichoke are t}-pical examples (Fig. 104-107) ; and the difference between these subterranean branches and the roots which they may bear is very obvious. Their eyes are axillary buds ; the leaves which subtend them are plainly dis- cernible, in the form of short and closely appressed scales. In the attempt, occasionally seen, to form axillary tubers above- ground by the Potato-plant, the leafy nature of the scales is evidenced. (Fig. 105.) By heaping the soil around the stems, the, number of tuber- 1 iferous branches may be in- creased. The number of nodes and internodes involved in a tuber may be many or few. There is one instance of what may in autumn, to be renewed by an axillary bud, which makes its subterranean growth and the rudiments of the aerial in early summer. 1 This rhizoma is a monopodium, being continued year by year by the terminal bud, and the aerial stem or stems sent up in spring, bearing the whorl of leaves and blossom, are axillary branches. FIG. 103. An older and longer one of the same species, showing branches, scars left by former leaf- and flower-bearing stems: also at tip (stripped of the covering scales), the bases of two such stems of the season, and the terminal bud between them, for the con- tinuation of the growth of the rootstock, &c., the next season. FIG. 104. Base of stem of Helianthus tuberosus, or Jerusalem Artichoke, developed from a tuber, and producing a second generation of tubers. FIG. 105. Monstrosity of a Potato-plant, with an axillary bud developing into some- thing between a bulblet and a tuber ; the scales represented by obvious leaves. (From the Gardeners' Chronicle.) (JO MORPHOLOGY OF STK.MS. be called u Monomerous tuber, namely in Nelumbium luteum (Fig. 108"), where it consists of a single thickened iuternode of 107 UK', an aquatic runner, which is accordingly quite destitute of scales or buds. The growth proceeding from this simple tuber is necessarily from a bud of the node at its apex, whence also a cluster of roots is produced. Of a somewhat similar nature are the con- catenate tubers of Apios tuberosa (several of which io8a are strung :is it were upon a long filiform axis), the tubers not uufreqtieiitly being mo- nomerous. although the larger ones are not so. 111". Tubwl«>s, as they may be termed, an- of a mixed or ambiguous character between tubers and tuberous roots. A good example of the latter is afforded by Dahlia-roots. (Fig. 68.) They yield their nourishing substance to growing buds on the stem above, but do not themselves normally produce even FIG. 10(5. Form ins |>nt:\t"i's in various stairs. 107. One of the younger ones en- larged, ins. Section of a small portion passing through an eye. or bud. more enlarged. FIG. 108a. A monomerous (i. e. one-membered) tuber of Nelumbium luteum, formed of a single internode. SPECIAL FORMS. 61 adventitious buds. Sweet potatoes (55), although equally roots, do produce adventitious buds, especially from near the upper end. The somewhat similar tubercles or tumefied roots of certain Orchises and other plants of the same tribe,1 definite in number and shape, and sometimes imitating a corm, are charged with a bud at the upper end, near their origin. Ap- parently, the origin is a bud from the base of the parent stem, which bud directly forms a tumefied short root from its' very base.2 118. A Corm (Cormus) is to be compared on the one hand with a short rootstock or tuber, on the other with a bulb. It is a subterranean fleshy stem, of rounded or depressed figure and solid texture. Some of its buds grow into new corms, and these, upon the death of or separation from the parent, become new individuals : some develop above ground , ^ n the vegetation and the blossoms of the season. A good type of corm is that of Cyclamen (Fig. 109), in which the very base of the seedling stem grows fleshy, and widens from year to year, but hardly at all lengthens, and so becomes far broader than high, or de- pressed. As the main bulk belongs to the 109 first internode, or caulicle, the buds from which the yearly growths of leaves and flower-stalks spring are at the centre of the summit or upper surface, the roots from the lower, and the sides seldom pro- duce any buds. The conn of Indian Turnip (Arisaema triphyUum, Fig. 110) is somewhat similar, but it sends up a single stout stem, and the roots spring" from around the base of this. These are completely naked corms. 119. But in Crocus (Fig. Ill, 112), Colchicum (Fig. 117),. Gladiolus, and the like, the sheathing bases of one or two leaves' enclose the corm with a membranous-scaly coat, giving it exactly the appearance externally of a coated bulb. Such have been not inappropriately named solid bulbs. In common parlance, they will doubtless continue to be called bulbs, and even in popular 1 Not, however, such as those of Aplectrura, Tipularia, etc., which are genuine corms or tubers. 2 Irmisch, Beitr. Biol. & Morphol. Orchid. 1853, fide Duchartre, 6le'm. Dot. 278. FIG. 109. Depressed corm of Cyclamen. FIG. 110. Corm of Indian Turnip, Arisaema triphyllum. 62 MORPHOLOGY OF STEMS. botanic.-il descriptions. In fact, while they differ from naked corms in having some investment, the}* differ from true bulbs only in the greater size of the solid axis and the fewness of /\ ^ the investing scales ; the stem or solid bod}* mnking the greater part of the corm, but a very small part of a proper bulb. There are, more- over, all gradations be- tween the two. A Bulb, as compared with a corm, may be said to be -an exceedingly abbreviated stem, reduced to a flat plate, from the lower face of which roots are produced, from the upper face, leaves in the form of scales ; these scales being either reduced and thickened leaves or the thickened bases of ordinary leaves. Compared with buds (73), it is a very flesh}- bud, usually large and subterranean, the axis of which never elongates. It is a 120. provision for future growth, the stored nourishment of which Is deposited in the leaves, or the homologues of leaves, instead of in the stem. FIG. 111. Corm of Crocus, the few thin envelopim; scales removed, showing their sears, \\hich mark the nodes, the shrivelled vestige of the last year's conn at the base, and lm. Is developing into new ones on various parts of its surface. iu. Vertical section of a similar corm, \\iili a terminal and one lateral luid. l''l( !• 1 !::. Section of a tnnieuted bulb of the Onion. FKi. II I Vertical section of the bulb of the Tulip, showing its stem or terminal budlc) and t \\o axillary buds (/». ft). FIG. 11"). Bull i of a Garlic, with a erop of young bulbs. KM;, in; Vertical section of the corm of a Crocus: 2), the cladophyU is wholly leaf-like in appearance as well as in function, and it never bears either scale-leaf or blossom ; but the flowers are on slender stalks from buds out of the same axil. (See Dickson in Trans. Bot. Soc. Edinb. xvi., and Van Tieghem, Bull. Bot. Soc. France, xxxi., for a discussion of the nature of cladophylla.) 128. To all such leaves or imitations of leaves. IJischoff has given the name PIIYLLOCLADIA, sing. PHYLLOI I.ADI KM. To those definitely restricted to one internode, and which so closely counterfeit leaves, Kunth gave the name of C I.ADODIA, sing. CLADODIUM. The best common name for all productions which imitate leaves would have been that of phyllodium (meaning simply a leaf-like body) ; but that term wras first applied and is restricted to the case of a petiole imitating the blade of a leaf. The name Phyllocladium (meaning a leaf-like branch) may properly be retained for the whole series of leaf-like bodies here de>cribed. But for those of the preceding paragraph, which are so peculiarly leaf-like, Knnth's name of Cladodium (i. e. a branch-like body) is false in meaning, and may be replaced by that of CLADOPIIYLUM (I.e. leaf- branch), or in shorter English C'LADOPIIYLL. 129. Frondose Steins. Finally, in some few phaenogamoua plants, tin- whole vegetation is re- duced ton, simple leaf-like expansion, as in Duckweed (Lemna), FIG. 122 Myrsiphylluni. with <-l:ihylls serving for foliage; the true leaves con- sisting of minute ami vrry Inconspicuous scales suMcnding tin1 former. I'" 1C. 123. Asinijr Hadojihyllof Ruscus aculeatus in the axil of a scale-leaf, bearing another scale-leaf on the middle of its face, and flowers in the axil of this. FIG. 124. l.i'iini.-i minor, a common Duckweed, whole plant in tlower, magnified. INTERNAL STRUCTURE. 67 Fif. 124. Here is no differentiation whatever into stem and O foliage ; but the expanded floating body which serves for both must be counted as stem developed horizontally into a flat plate, for it produces a root from the under surface and a flower from the edge. This simplification is common in some orders of Cryptogamous plants ; and such a bod}', which answers both for stem and foliage, is termed a FROND, from the Latin frons, which means either leaf or leafy bough. In some species of Lemna the frond is thickened or plano-convex : in Wolffia, the simplest and smallest of phaenogamous plants, it is a globular green mass, seldom much larger than the head of a pin, wholly destitute of root, propagated by proliferous budding from one side, and from within the top producing a flower or pair of flowers. § 3. INTERNAL STRUCTURE. 130. The investigation of the intimate structure of the stem, as of the other organs, belongs to vegetable anatomy or histology (treated in Part II.) ; but the general outlines of structure, so far as is requisite to the explanation of what is visible to the naked eye, should be here explained. 131. The stems of phsenogamous plants anatomically consist of two general elements, the cellular and the woody ; the former exemplified in the commoner stems by the pith and outer bark, the latter by the wood. Both are equally composed of cells, or origi- 124" nate as such ; but those which form the woody system of the stem mainly undergo, at a very early period, transformation into tubes, some of which are of such small calibre that their common name of fibres is not inappropriate ; others, of larger size or ampler calibre, take the name of ducts or vessels. The latter are almost FIG. 124a. A magnified slice of a portion of the flower-stalk of Richardia jEthiopica (the so-called Calla Lily), transverse with some longitudinal view : mainly parenchyma, the cells huilt up so as to leave comparatively large vacancies (intercellular spaces or air-passages) ; near the centre a cross-section of a fibro-vascular bundle, and next the margin or rind some liner ones. MOKPHOLOGY OF STEMS. always associated with the wood-cells, so that they are in a general way taken together as constituting the wood, or woody tissue, and as forming what is more definitely termed fibro- vascular tissue or, when distinguishable into threads, fibro-vascular bundles. These run lengthwise through the stem, sometimes as such separate threads, sometimes confluent into a compact structure. The softer or at least the non-fibrous portions, formed of comparatively a b d e fa A short and commonly thin-walled cells, form cellular tissue. Its ordinary form (of roundish, cubical, or polyhedral and thin-walled rells) is called parenchyma. This abounds in herbaceous stems or herbaceous parts: in trees and shrubs, woody tissue largely pre- vails; in most herbs, it forms a notable portion ; in some (especially FIG. 125. Fibro-vascular elements, a. Bast-cells (long wood-cells) of fibrous bark of Linden <>r p.ass-wood. It. Some wood-cells and (lirlowla duct, and r a detached wood-cell of the wood of same tree, equally magnified \\itli anana. /.'. Due! from Celery, the thread within spiral or annular In low, reti. nlated aliovc. and higher passing into the state of dotted duct. /. Duct from Impaticns, with Hi.' open spiral passing into rin;;s at the middle. All magnilied somewhat equally. INTERNAL STRUCTURE. 69 in certain aquatic herbs), it is reduced to a few threads or vessels, generally delicate, and sometimes obscure. The ac- companying anatomical illustrations (Fig. 124", 125, with their explanations) will give a general idea of the nature of the ana- tomical elements of the stem. K>2. In the forming state, the whole stem is parenclryma ; but an early differentiation takes place, converting certain portions into woody or fibro-vascular tissue. This is arranged in two ways, giving rise to two kinds of stem in phaenogamous plants, which have been termed the Endogenous and the Exogenous,1 meaning inside and outside growers. 133. The two plans of stem are usually manifested in external conformation as well as internal structure, and are correlated with important differences in embryo, foliage, and flower.2 Palms, Lilies, Rushes, and Grasses are examples of the endogenous class ; the ordinary trees and shrubs, especially those of cool climates, and a large part of the herbs, are of the exogenous class. In an exogenous stem, the wood occupies annual concentric layers, one of each }-ear's growth ; the centre is occupied by a pith, composed of parenchyma only, the circumference by a separable bark ; so that a cross-section presents a series of rings or circles of wood, or in the first year one ring, surrounding the pith and surrounded by the bark. An endogenous stem has the wood in distinct threads or fibro-vascular bundles, travers- ing the cellular system or parenchyma with little or no obvious order, and presenting on the cross-section the divided ends of these bundles in the form of dots ; these usually (but not always) diffused over 1 Terms introduced by DeCandoIle, following the ideas of Desfontaines, and which have played an important part in structural and systematic botany ever since DeCandoIle adopted these names as those of the two primary divisions of phasnogamous plants, Exogence and Endogencc. But it has long been seen that the name of the second kind is not appropriate ; and the older and better (though longer) names of Jussieu, Monocotyledones and Dicotyledones, are reverted to. Yet the Candollean names are still much employed, with due explanation, to designate the two kinds of structure of the stem. 1 Yet with some more or less valid exceptions, as when the annual stem of Podophyllum and the rhizoma of Nymphsca, among dicotyledonous plants, imitate the endogenous structure ; or where the pith of an evidently exogenous stem, as in the Piperaceaj, has scattered woody bundles in an endogenous fashion ; or where monocot3rledonous plants have all their woody bundles in a definite circle, as in Luzula, Croomia, &c. FIG. 126. Section of a small Palm-stem, in two directions. 70 MORPHOLOGY OF STEMS. the whole section, or when few in number of somewhat definite position or arrangement. The ordinary appearance of such a stem, both on the Longitudinal and the cross-section, is shown in Fig. 126 ; it may also be examined in the Cane or Rattan, the l»amboo, and in the annual stalk of Indian Corn or of Asparagus. The appearance of ordinary wood is very familiar. 135. The newer woody bundles of an endogenous stem are vari- ously intermingled with the old. When DeCandolle gave the name, it was supposed, from Desfontaines's researches, that the older bundles occupied, or came at length to occupy, the circumference of the trunk, while onby new ones were formed in the centre ; and that increase in diameter, when it took place at all, resulted from the gradual growth and distention of the whole. Hence the contrasting name of endogenous, or inside growing, and for such plants the name of ENDOGENOUS PLANTS, or ENDOGENS. Our actual knowledge of the structure and growth of these stems, as will be seen, cannot be harmonized with this view in any way which gives to the name endogenous an appropriate signifi- cation. The name continues as a counterpart to the more correct one of exogenous, and as a survival of former ideas. 136. The Endogenous Structure (so called) of the stem is cor- related with a monocotyledonous embryo (39), usually with a ternary arrangement in the flower (322), and commonly with parallel-veined leaves. (173.) Endogens. although they have many herbaceous and a few somewhat woody representatives in cool temperate climes, mostly attain their full variety of fea- tures and rise to noble arborescent forms under a tropical sun. Yet I 'a Iras --the arboreous type of the class — do extend as far north in this country as the coast of North Carolina (the natural limit of the Palmetto, Fig. 126") ; while in Europe the Date and the Chanuerops thrive in the warmest parts of the Euro- pean shore of the Mediterranean. The manner of their growth gives them a striking appearance ; their trunks being unbranehed cylindrical columns, rising to the height of from thirty to one hundred and fifty feet, and crowned at the summit with a simple cluster of peculiar foliage. Palms generally grow from the terminal bud alone, and perish if this bud be destroyed : they grow slowly, and bear their foliage in a cluster at the summit of the trunk, which consequently forms a simple cylindrical column. But in some instances two or more buds develop, and the stem branches, rarely and accidentally in ordinary specie's, regularly in the Doum Palm of Upper Egypt, and in the Pandanus, or Screw-Pine (Fig. 69), which belongs to a family allied to Palms : in such cases the branches are cylindrical. But when lateral INTERNAL STRUCTURE. 71 buds are freely developed (as in the Asparagus) , or the leaves are scattered along the stem or branches by the full development of internodes (as in m the Bamboo, Maize, &c. ) , the}' gradually taper upward in the manner of most ex- ogenous stems. 137. This kind of stem comprises several subordinate types as to internal structure, which to be well understood must be studied his- tologically, under the microscope.1 To one of these, by no means the simplest, belongs the ordinary palm- stem, the anatomy of which was made classical by Mohl, and has been supplemented by Naegx-li. In this a large part of the bundles, or all of the more conspicuous kind, starting from the base of the leaf to which they respectively belong, curve inward more or less strongly toward the centre of the stem, and thence gradually outward as they descend until they reach the rind, in which the attenuated lower extremity mostly terminates. Consequently, the bundles from different heights cross in their course, somewhat 1 For the best and most accessible memoir on the subject, of recent date, see Guillaud, Recherches sur 1'Anatomie comparee et le De'veloppement des Tissus de la Tige dans les Monocotyledones, published in Ann. Sci. Nat. ser. 6, v. 1-176, 1877. Six types of the stem of Monocotyledons are here recognized by anatomical characters and modes of growth, one of them having four modifications. FIG. 126°. Sabal Palmetto in various stages ; also the Yucca aloifolia or Spanish Bayonet. MORPHOLOGY OF STEMS. as shown in Fig. 127. It is partly owing to this connection of these fibres with the rind that the latter is not separable from the stem. In some Palms, and in Grasses, there is no marked distinction between the wood and rind, or no proper rind at all, In others, such as the Palmetto (Fig. 126), there is a marked rind or false bark, which receives independent fibro-vascular bundles from the leaf-stalk-, and is travelled by them in parallel lines. In Grass-stems, and others with loim- inter- O nodes and closed nodes, the fibro-vascular bundles all run approximately straight and parallel through the internodes. but are intricate and anastomosed in the nodes. The whole centre of the ink-modes, when not hollow or before it becomes so, is occupied by a true pith, like that of an Exogen, and in some cases equally destitute of fibro-vascular bundles, but often with scattered ones, after the manner of certain Exogens anomalous in this respect, such as Nyctaginacese and some Araliacea-. Endogenous stems of simpler structure, as in herbaceous Liliaeea, Commelynacese, &c., have a distinct cortical portion (at least in the root-stock or portion of stem properly comparable with palm-trunks and the like) ; but this is mostly destitute of fibro-vascular bundles. Most of them have two kinds of vascular bundles, one of which not rarely occupies an exact circle in the line of division between the cortical and medullary portion (between bark and pith), and the other is within this circle, either of very few and scattered bundles, as in Convallaria majalis. or numerous and scattered, as in 1'vularia and the leafy stems of Tradescantia Virginica ; or these bundles are few and arranged nearly in an inner circle close around the centre. Finally, Luzula and Croomia have only one kind of bundles, answering to the outer ones of Convallaria ; in other words, the woody system forms a simple circle, dividing a purely cellular medullary from a similar cortical portion, thus closely imitating an herbaceous exogenous stem of the same age. l.'irt. An annual endogenous stem increases in diameter by general growth until it attains its limit. Ligneous and enduring stems increase similarly up to u certain period. Then the rind FIG. \'il. Diagrammatic view of the curved course of the fibro-vascular bundles in a palm-trunk. INTERNAL STRUCTURE. 73 sooner or later ceases to distend or adapt itself to further in- crease in diameter, and there is no interior provision for indefinite increase in the greater number of woody endogenous trunks. But in Dracaena (Dragon-trees) , in the arborescent Yuccas, and the like, the zone intermediate between the cortical and interior re- gion, which is for a tune active in many Endogens, here grows continuously and indefinitely. Such trunks increase in diameter throughout life ; they may attain a very great age (as some Dragon-trees have done) ; and they imitate exogenous trunks to a considerable extent in mode of growth. 139. The wood of an endogenous woody stem is hardest and most compact at the circumference ; in palm-stems commonly it is largely mixed with parenchyma or pith at the centre, even in old trunks. 140. The Exogenous Structure, that of ordinary wood, is char- acterized by the formation of a distinct zone of wood between a central cellular medullary portion (pith) and an outer chiefly cellular portion (bark) , traversed by plates from the pith (medul- lary rays) , and b}- increasing from the outer surface of this zone between wood and bark, the increase in enduring stems consist- ing of definite concentric annual layers. 141. Its Beginning, at the earliest growth of the embryo, is in the appearance of a few ducts (Fig. 125,/-/), at definite points in the common parenchyma of the initial stem (four equidistant ones in the Sugar Maple) ; each is soon surrounded by incipient proper wood-cells (Fig. 125, 5, c), together forming a fibro-vascular bundle or thread. Additional ones are intercalated as the second and third internodes develop, and so a column (in cross- section a ring) of wood is produced, always so arranged as to FIG. 128. Diagram of a cross-section of a forming seedling stem, showing the manner in which the young wood is arranged in the cellular system. FIG. 129. The same at a later period, the woody bundles increased so as nearly to fill the circle. FIG. 130. The same at the close of the season, where the wood lias formed a com- plete circle, interrupted only by the medullary rays, which radiate from the pith to the bark. 74 MORPHOLOGY OF STEMS. 132 surround a purelj' cellular central part (the pith) , while sur- rounded b_v a cellular external rind, the bark, or outer bark. The diagrams (Fig. 128-130) rudely show some stages in trie formation of the zone of wood. The fibro- vascular bundles originate in the bases of the leaves, and develop outward into the forming leaves as well as downward into the forming stem. 1 12. First Tear's Growth. The wood, even in a herbaceous or annual stem, at the completion of the first year's growth, forms a zone or tube, enclosing the pith. But it is traversed by plates (in cross-section lines) of parenchyma, or eel lular tissue of the same nature as the pith, which radiate from that to the bark, and thus divide the wood into wedges. These lines, forming what is called the silver-grain in wood, are the MED- ULLARY RAYS. They rep re si -I it the cellular system of the wood it- sell', or untransformed parenchyma. Being pressed by the woody wedges, their cells are laterally flattened. In some stems, the med- ullary rays, or many of them, are comparatively broad and conspicuous; in others. Him and inconspicuous or irregular. The growth of the woody wedges is soon complete, except at the outer portion, next the bark : here they usually continue to grow through the season : that, is the wood grows externally. The general ana- Kl<;. l.'il. Longitudinal and i i-.-ms\ .•]>,• section of a stem of the Soft Maple ( Acer dasyearpuni). at tin- rlnsi> of the lirst year's growth; of the natural size. FNS. l.'S'J. Portion of tin same, magnified, showing the cellular jiitli, surrounded by the wood, and that l>y the bark. FIG. 133. More inajrniiied slice of the same, reaching from the bark to the pith: a. part of the pith ; b. vessels of the medullary sheath ; c. the wood ; f tin- iiii'ilulliir.v rays, passing transversely from the pith (/)) to the bark(/;): magnified. But. a section can seldom be made so as to show one unbroken plate stretching across the wood, as in this instance. FIG. 136. A vertical section across the ends of the medullary rays: magnified. INTERNAL STRUCTURE. 77 stem, and of the whole plant. Lastly, the inner bark, accord- ingly named ENDOPHLGEUM, takes the special name of LIBER, and is the most important portion of the bark in the stems of trees and shrubs. Complete and well-developed liber, like that of Linden or Bass wood, contains two peculiar kinds of cells in InWUffMns&i^VHDHQai M'^oQ^mHHW^m 135° addition to common parenchyma, both of the fibrous or vascular class: viz., 1. CRIBRIFORM or SIEVE-CELLS, a sort of ducts the walls of which have open slits, through which they communicate with each other; 2. BAST or BAST-CELLS, the fibre-like cells which give to the kinds of inner bark that largely contain tlu'in FIG. 135a Portion of a transverse section (above), and a corresponding vertical sec- tion (below), magnified, reaching from the pith ip) to the epidermis (e) of a stem of Negundo, a year old : B. the bark ; W. the wood ; and C. the cambium-layer, as found in February. The parts referred to by small letters are: p. a portion of the pith: mr. small portion of a medullary ray where it runs into the pith; four complete med- ullary rays as seen on a transverse section, appear in the upper figure, running from pith to bark : ms. medullary sheath, a circle of spiral unreliable ducts, one seen length- wise with uncoiling extremity in the lower figure: w, w. woody tissue: (Id. one of the dotted ducts interspersed in the wood : cl. cambium-layer or zone of new growth of wood and inner bark : l-b. liber or inner bark, the inner portion of which is here cellular, the outer (6) composed of slender and thick-walled bast-cells or true liber-cells: ge. green envelope or inner cellular bark: ce. corky envelope or outer cellular bark: e. epidermis. 78 MORPHOLOGY OF STEMS. their strength and toughness. They are like wood-cells except in their greater length and flexibility, and in the thickness of their walls, which great ry exceeds the calibre. This is the material which gives l<> the bast or inner bark of Basswood, &c., the strength and pliability that adapts it for cordage and for making mats: it is the material of linen, and the like textile fibres. ( For a view of the whole composition and structure of a woody stem at the close of the first 3-ear's growth, and immedi- ately before that of the second year begins, see Fig. 135". ) 14-'). Annual Increase in Diameter. An herbaceous stem does not essentially differ from a woody one of the same age, except that the1 wood forms a less compact or thinner zone; and the whole perishes, at least down to the ground, at the close of the season. But a woody stem makes provision for continuing its growth from year to year. As the layer of wood continues to increase in thickness throughout the season, b}* the multiplication of cells on its outer surface, between it and the bark, and when growth ceases this process of cell-multiplication is merely sus- pended, so there is always a zone of delicate young cells in- terposed between the wood and the bark. This is called the CAMBIUM, or, better, the CAMBIUM-LATER. It is charged with organizable matter, which is particularly abundant and mucila- ginous in spring when growth recommences. This mucilaginous matter was named Con//// inn by the older botanists: they sup- posed— as is still popularly thought-- that the bark, then so readily separable, really separated from the wood in spring, that a quantity of rich mucilaginous sap was poured out between them and became organized into a tissue, the inner part becoming new wood, the outer, new bark. But delicate slices show that there is then no more interruption of the wood and inner bark than at any other season. The bark, indeed, is then very readily detached from the wood, because the cambium-layer is gorged with sap; but such separation is effected by the rending of a delicate forming tissue. And if some of this apparent mucilage be scraped oil' from the surface of the wood, and examined under a good microscope, it will be seen to be a thin stratum of young wood-cells, with the ends of medullary rays here and there in- terspersed. The inner portion of the cambium-layer is therefore nascent wood, and the outer is nascent bark. As the cells of this layer multiply, the greater number lengthen vertically into woody tissue: some are transformed into ducts; and others, remaining as parenchyma, continue the medullary rays or com- mence new ones. In this way. a second layer of wood is formed the second season over the whole surface of the former layer INTERNAL STRUCTURE. 79 between it and the bark ; and this is continuous with the woody layer of the new roots below and of the leafy shoots of the sea- son above. Each succeeding year another layer is added to the wood in the same manner, coincident with the growth in length by the development of the buds. A cross-section of an exoge- nous stem, therefore, exhibits the wood disposed in concentric rings between the bark and the pith ; the oldest lying next the latter, and the youngest occupying the circumference. Each layer being the product of a single year's growth, the age of an exogenous tree may, in general, be correctly ascertained by counting the rings in a cross-section of the trunk.1 144. Demarcation of the Annual Layers results from two or more causes, separate or combined. In oak and chestnut wood, and the like, the layers are strongly defined by reason of the accumu- lation of the large dotted ducts (here of extreme size and in great abundance) in the inner portion of each layer, where their open mouths on the cross-section are conspicuous to the naked e}Te, making a strong contrast between the inner porous and the exterior solid part of the successive layers. In maple and beech wood, however, the ducts are smaller, and are dispersed through- out the whole breadth of the layer ; and in coniferous wood, viz. that of Pine, Cypress, &c., there are no ducts at all, but only a uniform wood}' tissue of a peculiar sort. In all these, the de- marcation between two layers is owing to the greater fineness of the wood-cells formed at the close of the season, viz. those at the outer border of the layer, while the next layer begins, in its 1 The annual layers are most distinct in trees of temperate climates like ours, where there is a prolonged period of total repose, from the winter's cold, followed by a vigorous resumption of vegetation in spring. In tropical trees, they are rarely so well defined ; but even in these there is generally a more or less marked annual suspension of vegetation, occurring, however, in the dry and hotter, rather than in the cooler season. There are numerous cases, moreover, in which the wood forms a uniform stratum, whatever be the age of the trunk, as in the arborescent species of Cactus ; or where the layers are few and by no means corresponding with the age of the trunk, as in the Cycas. In many woody climbing or twining stems, such as those of Clematis, Aristolochia Sipho, and Menispermum Canadense, the annual layers are rather obscurely marked, while the medullary rays are unusually broad ; and the wood, therefore, forms a series of separable wedges disposed in a circle around the pith. In the stem of Bignonia capreolata, the annual rings, after the first four or five, are interrupted in four places, and here as many broad plates of cellular tissue, belonging properly to the bark, are inter- posed, passing at right angles to each other from the circumference towards the centre, so that the transverse section of the wood nearly resembles a Maltese cross. But these are exceptional cases, which scarcely require notice in a general view. 80 MORPHOLOGY OF STEMS. vigorous vernal growth, with much larger colls, thus marking :m abrupt transition from one layer to the next. Besides being finer, the later wood-cells of the season are commonly flattened antero-posteriorly, probably by growing under greater pressure. 14;">. Kadi layer of wood, once formed, remains essentially iinch.-iiiged in position and dimensions. But, in trunks of con- siderable age, the older layers undergo more or less change in color, density, perviousness to moisture, &c. Mil. Sap-wood (ALBruxuM). In the plantlet and in the developing hud, the sap ascends through the whole tissue, of whatever sort : at first through the parenchyma, for there is then no other tissue ; and the transmission is continued through it, especially through its central portion, or the pith, in the growing apex of the stem throughout. But, in the older parts below, the pith, soon drained of sap. becomes filled with air in its place, and thenceforth it hears no part in the plant's nourishment. As soon as wood-cells and ducts are formed, they take an active- part in the conveyance of sap, for which their tubular and ca- pillary character is especially adapted. But, the ducts in older parts, except when gorged with sap, contain air alone ; and in woocty trunks the sap continues to rise year after j-ear to the places where growth is going on, mainly through the proper woody tissue of the wood. In this transmission, the new layers are most active ; and these are in direct communication with the new roots on the one hand and with the buds or shoots and leaves of the season on the other. So, by the formation of new annual la}-ers outside of them, the older ones are each year removed a step farther from the region of growth ; or rather the growing stratum, which connects tin- fresh rootlets that imbibe with the foliage that elaborates the sap. is each year removed farther from them. The latter, therefore, after a lew years, cease to convey sap, as they have long before ceased to take part in any vital operations. The cells of the older layers, also, usually come to have thicker walls and smaller calibre than those of the newer. Thus arises a distinction — sometimes obscurely marked, some- times :dirnpl and conspicuous — into sap-wood and /irtir'-wood. The former is the popular name given to the outer and newer layers of -.i>|'|er. more open, and bibulous wood. The early physi- ologists named it . While the newer layers of the wood abound in crude sap. which they convey to the leaves, those of the inner bark abound in elaborated sap. which they receive from the leaves and comey to the cambium-layer or zone of growth. The proper juices and peculiar products of plants are according!}' found in the foliage and the bark, especially in the latter. In the bark, therefore (either of the stem or of the root) , medicinal and other principles are usually to be sought, rather than in the wood. Nevertheless, as the wood is kept in connection with the bark by the medullary rays, many products which probably originate in the former are deposited in the wood. INTERNAL STRUCTURE. 83 153. The Living Parts of a tree or shrub, of the exogenous kind, are obviously only these: 1st, The summit of the stem and branches, with the buds which continue them upwards and annually develop the foliage. 2d, The fresh roots and rootlets annually developed at the opposite extremity. 3d. The newest strata of wood and bark, and especially the interposed cambium- la}'er, which, annually renewed, maintain a living communication between the rootlets on the one hand and the buds and foliage on the other, however distant they at length may be. These are all that is concerned in the life and growth of the tree ; and these are annually renewed. The branches of each year's growth are, therefore, kept in fresh communication, by means of the newer layers of wood, with the fresh rootlets, which are alone active in absorbing the crude food of the plant from the soil. The fluid they absorb is thus conveyed directly to the branches of the sea- son, which develop leaves to digest it. And the sap they receive, having been elaborated and converted into organic nourishing matter, is partly expended in the upward growth of new branches, and partly in the formation of a new layer of wood, reaching from the highest leaves to the remotest rootlets. 154. Longevity of trees. As the exogenous tree, therefore, annually renews its buds and leaves, its wood, bark, and roots, — every thing, indeed, that is concerned in its life and growth, - there seems to be no necessary cause, inherent in the tree itself, why it may not live indefinitely. Some trees are known to have lived for one and two thousand }rears, and some are possibly older.1 Equally long may survive such endogenous trees as the Dragon tree (Dracaena) , which have provision for indefinite in- crease in diameter (138), and for the production of branches. The famous Dragon tree of Orotava, in TenerifFe, now destroyed by hurricanes and other accidents, had probably reached the age of more than two thousand years. 155. On the other hand, increase in height, spread of branches and length of root, and extension of the surface over which the annual layer is spread, are attended with inevitable disadvantage, which must in time terminate the existence of the tree in a w:iy quite analogous to the death of aged individual animals, which is not directly from old age, but from casualties or attacks to 1 The subject of the longevity of trees has been discussed by DeCandolle, in the "Bibliotheque Universelle" of Geneva, for May, 1831, and in the second volume of his " Physiologic Ve'ge'tale ; " more recently, by Alphonse DeCan- dolle in the " Bibliotheque Universelle ; " and in this country by myself in the " North American Review," for July, 1844. For an account of the huge Red- woods (Sequoias) of California, see Whitney's Yosemite Book. 84 MORPHOLOGY OF STEMS. which the aged are either increasingly incident or less able to resist. A tree like the Banyan (/>!», Fig. 71), which by aerial roots continues to form new trunks for the support and sustenance of the spreading brandies, and thus ever advances into new soil, has a truly indefinite existence ; but, then, it becomes a forest, or is to be likened to a colony propagated and indefinitely in- creased by suckers, offsets, or other subterranean shoots. So the question of the secular continuation of the individual plant becomes merged in that of continuation of the race. — at least of a bud-propagated race, — the answer to which is wholly in the domain of conjecture.1 However this ma'y be, it is evident that :i vegetable of the higher grade is not justly to be compared with an animal of higher grade ; that individuality is incompletely realized in the vegetable kingdom;2 that rather 156. The Plant is a Composite Being, or Community, lasting, in the case of a tree, through an indefinite and often immense num- ber of generations. These are successively produced, enjoy a term of existence, and perish in their turn. Life passes onward continually from the older to the newer parts, and death follows, with equal step, at a narrow interval. Xo portion of the tree is now living that was alive a few years ago ; the leaves die annu- ally and are cast off, while the internodes or joints of stem that bore them, as to their wood at least, buried deep in the trunk under the wood of succeeding generations, are converted into lifeless heart-wood, or perchance decayed, and the bark that belonged to them is thrown off from the surface. It is the aggre- gate, the blended mass alone, that long survives. Plants of single cells, and of a definite form, alone exhibit complete indi- viduality ; and their existence is extremely brief. The more complex vegetable of a higher grade is not to be compared with the animal of the highest organization, where the oll'spring always separates from the parent, and the individual is simple and indi- visible. But it is truly similar to the branching or arborescent coral, or to other compound animals of the lowest grade, where successive generations, though capable of living independently and sometimes separating spontaneously, yet are usually devel- oped in connection, blended in a general body, and nourished more or less in common. Thus, the coral structure is built up by the combined labors of a vast number of individuals, — by the successive labors of man}' generations. The surface or the recent shoots only are alive ; beneath are only the dead remains 1 See Darwiniana, xii. 338-355. 2 As, perhaps, was first explicitly stated by Engelmann, in his inaugural essay, De Antholysi Prodromus, Introduction, § 4. MOKPHOLOGY OF LEAVES. 85 of ancestral generations. As in a genealogical tree, only the later ramifications are among the living. The tree differs from the coral structure in that, as it ordinarily imbibes its nourish- ment mainly from the soil through its roots, it makes a downward growth also, and, by constant renewal of fresh tissues, maintains the communication between the two growing extremities, the buds and the rootlets. Otherwise, the analogy of the two, as to individuality, is well-nigh complete. SECTION IV. OF LEAVES. § 1. THEIR NATURE AND OFFICE. 157. Leaf (Lat. Folium, in Greek form Phyllum), as a botani- cal term, has on the one hand a comprehensive, on the other a restricted sense. In its commonest sense, as used in descriptive botany, it denotes the green blade only. Yet it is perfectly understood that the footstalk is a part of the leaf, and therefore that the phrase " leaves cordate," or the like, is a short wa}' of saying that the blade of the leaf is cordate or heart-shaped. Moreover, two appendages, one on each side of the base of the footstalk, when there is any, are of so common occurrence that they are ranked as a proper part of the organ. So that, to the botanist, a typical leaf consists of three parts : 1, BLADE or LAMINA ; 2, FOOT-STALK or LEAF-STALK, technically PETIOLE ; 3, A pair of STIPULES. (Fig. 142.) 158. The blade, being the most important parf of an ordinary leaf, may naturally be spoken of as the whole. Petiole and stipules are indeed subsidiary when present, and are not rarely wanting. Yet sometimes they usurp the whole function of foli- age, and sometimes there is no such distinction of parts. 159. Physiologically, leaves are green expansions borne by the stem, outspread in the air and light, in which assimilation (3) and the processes connected with it are carried on. Vegetable assimilation, — the most essential function of plants, being the conversion of inorganic into organic matter, — takes place in all ordinary vegetation only in green parts, and in these when exposed to the light of the sun. And foliage is an adaptation for largely increasing the green surface. But stems, when green, take part in this office in proportion to the amount of surface, sometimes monopolize it, and in various cases increase their means of doing so by assuming leaf-like forms. (126-129.) Leaves, especially in such cases, may lose this function, appear only as useless vestiges, or may be subservient to various wholly 86 MORPHOLOGY OF LEAVES. different uses. Form and function, therefore, are not sure indi- cations of tlu- trur nature of organs. 1GU. Morphologically, and in the most comprehensive sense, leaves are special lateral outgrowths from the stem, definitely and symmetrically arranged upon it; in ordinary vegetation and in the most general form constituting the assimilating apparatus (or foliage), but also occurring in other forms and subserving various uses. Sometimes these uses are combined with or sub- sidiary to the general function of foliage ; sometimes the leaf is adapted to special uses only. So the botanist — recognizing the essential identity of organs, whatever their form, which appear in the position and conform to the arrangement of leaves - discerns the leaf in the cotyledons of a bean or acorn, the scale of a lily-bull) or the coat of an onion, the scale of a winter bud, and the petal of a blossom. Therefore, while expanded green leaves (which may be tautologically termed foliage-leaves) are taken as the proper type, the common name of leaves, in the lack of any available generic word, is in morphological language extended to these special forms whenever it becomes needful to express their uhylline or foliar nature. 161. In the morphological view, all the plant's organs except- ing roots (and excepting mere superficial productions, such as hairs, prickles, &c.), belong either to stem or to leaves, are either cauline or phylline in nature. To the latter belong all the primary outgrowths from nodes, all lateral productions which are not axillary.1 Whatever is produced in the axil of a leaf is cauline, and when developed is a branch. 1(12. The Duration of Leaves is transient, compared with that of the stem. They may be fugacious, "when they fall off soon after their appearance ; deciduous, when they last only for a single season ; and persistent, when they remain through the cold season, or other interval during which vegetation is interrupted, and until after the appearance of new leaves, so that the stem is never lealless. as in Evergreens. In many evergreens, the leaves have only an annual duration ; the old leaves falling soon after those of the ensuing season are expanded, or. if they remain longer, ceasing to bear any active part in the economy of the vegetable, and soon losing their vitality altogether. In Pines and Firs, however, although there is an annual fall of leaves either in autumn or spring, yet these were the produce of some 1 Tlu'tv :ire cases in which this rule is of difficult application, or is seem- ingly violated, sometimes by the suppression of the subtending leal, as in the inflorescence of Crucifcne, rarely in other ways, to be explained in the proper places. THEIR STRUCTURE AND FORMS. 87 season earlier than the last ; and the branches are continually clothed with the foliage of from two to five, or even ten or more successive years. On the other hand, it is seldom that all the leaves of an herb endure through the whole growing season, the earlier foliage near the base of the stem perishing while fresh leaves are still appearing above. In our deciduous trees and shrubs, however, the leaves of the season are mostly developed within a short period, and the}T all perish in autumn nearly simultaneously. 163. Leaves soon complete their growth, and have no power of further increase. Being organs for transpiration, a veiy large part of the water imbibed b}- the roots is given out ~by the foliage, leaving dissolved earthy matters behind. Assimilation can take place only in fresh and vitally active tissue. It is incident to all this that leaves should be of only transient duration, at least in their active condition. 164. Defoliation. The leaves of most Dicotyledons and some Monocotyledons separate from the stem and fall by means of an articulation at the junction with the stem, which begins to form early in the season and is completed at the close. There is a kind of disintegration of a transverse layer of cells, which cuts off the petiole by a regular line, and leaves a clean scar, such as is seen in Fig. 81, 85, 91. Some leaves, notably those of Palm?, Yucca, and other endogens, die and wither on the stem, or wear away without falling. 165. In temperate climates, defoliation mostly takes place at the approach of winter. In warmer climates having only winter rain, this occurs in the hot and dry season. 166. Normal Direction or Position. The leaf-blade is expanded horizontally, that is, has an upper and an under surface. When erect, the upper surface faces the axis which bears it. To this, there are many seeming but no real exceptions ; that is, none which are not explicable as deviations or changes from the normal condition. (213-217.) § 2. THEIR STRUCTURE AND FORMS AS ORGANS OF ASSIMILATION OR VEGETATION, i. e. AS FOLIAGE. 167. The Internal Structure or Anatomy of the leaf needs here to be examined so far as respects its obvious parts and their general composition. The leaf, like the stem, is composed of two elements (131), the cellular and the woody. The cellular portion is the green pulp or parenchyma, and in this the work of assimilation is carried on. The woody is the fibrous frame- 88 MORPHOLOGY OF LEAVES. work, the separate parts or ramifications of which form what are variously culled tin- /•//«, -- a sufficiently proper term, — nert-es or veins. The latter names may suggest false analogies ; but they are of the commonest use in descriptive botany. That of reins. and of its diminutive, /v///Av.\-, for the smaller ramifications, is not amiss; for the fibrous framework not only gives firmness and support to the softer cellular apparatus. /. f thr epidermis of the ('..-inlcn Kalsam, with three atomata (all IT Hroiiguiart ). FIG. ].'«». Mii^niliril view of the 10,000th part of a square inch of the epidermis of the lower surface »f the leaf of the White Lily, with its stomata. 140. A single stoma, ii ina^nitieii. 141. Another stoma, widely open. THEIR STRUCTURE AND FORMS. 91 which have exogenous stems and at least dicotyledonous embiyos. The latter prevails in ordinary plants with exogenous stem and dicotyledonous embryo. 173. Parallel-veined or Nerved leaves (of which Fig. 143 is an illustration) have a framework of simple ribs (called by the earlier botanists nerves, a name still used in descriptions), which run from the base to tip, or sometimes from a central strong rib to margin of the leaf, in a generally parallel and undivided way, and sending off or connected by minute veinlets only. Grasses, Lily of the Valley, and the like, illustrate the commoner mode in which the threads of wood run from base to apex. The Banana and Canna are familiar illustrations of a mode not un- common in tropical or subtropical endogens, in which the threads or "nerves" run from a central rib (midrib} to the margin. Parallel- veined leaves are generally entire, or at least their margins not toothed or indented. The principal exception to this occurs when the ribs or the stronger ones are few in number and radiately divergent, as in the fiabeUiform leaves of Fan- palms, a peculiar modification of the parallel- veined type. Between leaves with nerves wholly of basal origin, and those with nerves all springing from a midrib, there are various grada- tions, and also in respect to curving. But parallel- veined or nerved leaves may be classified into FIG. 142. A leaf of the Quince, of the netted-veined or reticulated sort, with blade (6), petiole or leaf-stalk (/»), and stipules (st). FIG. 143. Parallel-veined leaf of the Lily of the Valley, Convallaria majalis. 92 .MORPHOLOGY OF LEAVES. . that is, with the nerves all springing from the base of the leaf, and Costal-nerrt'i/. springing from a midrib or costa. Either may be Rectinerved, the nerves running straight from origin to apex or margin of the leaf, as the case may be ; Ourn'itrrred, when curving in their course, as in the leaves of Funkia and in Canna ; Flabellinerved, where straight nerves and ribs radiate from the apex of the petiole, as in Fan-palms and the Gingko tree. 174. In typical parallel-veined leaves, all reticulation is con- lined to minute and straight cross-veinlets : in man}", these are coarser, branching, and reticulated ; in some, as in Smilax and Dioscorea, only the primary ribs or strongest nerves are on the parallel-veined plan ; the space between being filled with reticu- lations of various strength ; thus passing by gradations into 175. Reticulated or Netted- veined leaves. In most of these, from one to several primary portions of the framework are particularly robust, and give origin to much more slender ram- ifications, these to other still smaller ones, and so on. The strong primary portions are RIBS (costce) ; the leading ramifi- cations, VEINS (vence) ; the smaller and the ultimate subdivisions, VK INLETS (yenidce). All or some of the veins and veinlets are said to anastomose, i. e. variously to connect with those from other trunks or ribs, apparently in the manner of the veins and arteries of animals, forming meshes. But, as there is no opening of calibre of one into another, the word is etymologic-ally rather misleading. More properly, it is said that the veins or veinlets form reticulations or net-work. A primary division of rwticuluted leaves, and indeed of nerved leaves also, into two classes, is founded upon the number of primary ribs. 17G. There may be only a single primary rib; this traversing the blade from base to tip through its centre or axis (as in Fig. 142, 1.V2-156) is called the MIDHIB. There may be others, gen- erally few (one, two. three, or rarely four), rising from the apex of the petiole on each side of the midrib, running somewhat par- allel with it or more or less diverging from it: these are lateral ribs. Among parallel-veined leaves, the Banana, Canna. &c., have a single rib, from which the veins (in the older nomencla- ture here called nerves) all proceed. Most Lilies and the like have several approximately parallel ribs, but the midrib pre- dominant: in other cases, the midrib is no stronger than the others. In Fan-palms, the ribs are radial el y divergent, giving a tan-shaped or rounded outline to the blade. In reticulated leaves, in which the veins all spring from the ribs, the two THEIR STRUCTURE AND FORMS. 93 classes into which they divide are the pinnately veined and the palmately veined. 111. Pinnately or Feather- veined (or Penninerved) leaves are accordingly those of which the veins and their subdivisions are side branches of a single central rib (midrib), which traverses the blade from base to apex ; the veins thus being disposed in the manner of the plume on the shaft of a feather. (Fig. 142, 152, &c.) Sometimes these continue straight and undiminished from midrib to margins (straight- veined, as in Beech and Chest- nut, Fig. 152), sending off only small lateral veinlets ; some- times they ramify in their course into secondary or tertiary veins, and these into veinlets. Pinnate venation in reticulated leaves naturally belongs to leaves which are decidedly longer than wide. 178. Some of the primary veins, commonly among the lower, may be stronger than the rest, and thus take on the character of ribs, or by gradations pass into such. The leaf of the common annual Sunflower (Fig. 155) becomes in this way triple-ribbed or tripli-nerved. The appearance of a second pair of such strength ened veins makes the venation quintupli-ribbed o? quintupli-nerved. 144 147 148 149 157 Through the approximation of such strong veins to the base of the blade, this venation may pass into the 179. Palmately, Digitately, or Radiately Veined (or Palmi- nerved) class, of which leaves of common Maples and the Vine are familiar examples. (Also Fig. 158-160, &c.) In these FIG. 144-157. Various forms of simple leaves, explained in the text and in the Glossary. 94 MORPHOLOGY OF LEAVES. there are three, five, seven, or sometimes more ribs of equal strength, the central being the midrili. ;m. 151. Also Fig. 149, Pontcderia, a leaf of the parallel- veined class. 11' a if« mi. or Kidney-shaped, ]ike the last, only rounder and broader than long. (Fig. 158.) Aurli-iiluh', or Eared, having a pair of small and blunt pro- jections, or ears, at the base, as in Magnolia Fraseri. Fig. 17s. Sayittate, or Arrow-shaped, where such ears are acute and turned downwards, while the main body of the blade tapers upwards to a point, as in the com- mon species of Sagit- taria or Arrow-head, and in the Arrow- leaved Polygonum. (Fig. 165, 177.) Hastate, or Jlalberd- slntped, when such lobes at the base point, out wards, giving, the leaf the shape of the halberd of the olden time, as in Polygonum arifolium (Fig. 17l>) and Sorrel, Fig. 1G3. 182. Peltate or Shield-shaped leaves are those in which a blade of rounded or sometimes of other shape is attached to the petiole by some part of the lower surface, instead of the basal margin : those of Water-shield or Ilrasenia, of Nelumbium. and of Ilydro- cotyle umbellata are marked examples. The anomaly is mor- phologically explained b\ a comparison with deeply cordate or remlbrm leaves having a narrow sinus, such as those of Nym- pluca or Water Lily, and by supposing a union of the approxi- mated edges of the sinus. Fig. I.V.I and 1 ('»<>, from two species of Ilydrocotyle. one with open and the other with dosed sinus obliterated by the union, illustrate this. is:1,. As to llxtn'init.v, whether base or apex, there are several descriptive terms, expressive of the principal modifications; such as Acuminate, tapering, either gradually or abruptly, into a narrow more or less prolonged termination. (Fig. 180.) 179 FIG. 177-179. Sagittate, auriculate, ami hastate leaves. THEIR STRUCTURE AND FORMS. 97 Acute, ending in an acute angle, without special tapering, as in Fig. 181. Obtuse, ending with a blunt or roundish extremity, Fig. 182. Truncate, with termination as if cut off by a straight transverse line, as in Fig. 183. Refuse, with an obtuse extremity slightly depressed or re-enter- ing, as in Fig. 184. Emarginate, with a more decided terminal notch, Fig. 185. Obcordate, inversely heart-shaped, i. e. h'ke cordate, but the broader end and its strong notch at apex instead of base, Fig. 186. This and the following terms are applicable to apex only. Mucronate, abruptly tipped with a small and short point, like a projection of the midrib, as in Fig. 18f lobes, when definitely marked, may come into the descriptive phrase, as pinnately 7-lobed, pinnately 1 -cleft, parted, or divided, as the case may be. 190. Similarly, Figures 199 to 202 represent, respectively, a palmately three-lobed, three-cleft, three-parted, and three-divided, or, in Latin form, trilobate, trifid, tripartite, and trisect or trisected leaf. Fig. 166 is a palmately ^-parted leaf; Fig. 164, palmately FIG. 195-198. Pinnatcly lobed, cleft, parted, and divided leaves. FIG. 199-202. Palmately 3-lobed, cleft, parted, and divided leaves. 100 MORPHOLOGY OF LEAVES. ^c. Fig. H;2, a leaf of Dragon Arum, is palmntch/ :i-/»//W. But, as the lateral sinuses are not so deep as the others, the leaf is said to be pedately parta/, or pi-date, in the early terminology. P.M. .Moreover, as the lobes or divisions of a leaf mav be again similarly lolied or parted, &c., this composition may be indicated by the prefix twice, thrice, &c., as twice )>i,i,,,itif,d or bipinnatifid, llu-i<-<- /i/nnately parted, thrice palmateh/ parted, and the like. Thus, a word or two, or a short phrase, may describe even a complex leaf, so as to convey a perfectly clear and defi- nite idea of its conformation. 192. A distinction should now be drawn between simple and compound leaves. The distinction cannot be both natural and absolute ; lor the one may pass variously into the other. .SV/////A- leaves, which have been thus far considered, have a single lamina or blade, which may, however, at one extreme be entire, at the other many-parted, and even several times divided. l'.i:>. Compound Leaves are those which have- from two to many distinct blades, on a common leafstalk. These blades, called LEAKI.KTS, may be sessile on the common leafstalk, or thev may have leafstalks of their own. As the leaf very commonly separates in age by an articulation of its petiole with the stem, so leaflets are commonly more or less articulated with the com- mon petiole. When the leaf, with its petiole, falls from the stem, the leaflets may as completely separate from the common petiole. They do not always do this. Divided Leaves, such as those of Fig. 198 and 202, though ranked among the simple sorts, are compound in the sense of having distinct blades, but without articulation. Some of these blades are apt to be confluent; that is, a divided leaf is often in part merelv parted, as in the upper portion of Fig. l!)S. Such leaves are so inter- mediate between simple and compound that it becomes indilfer- enl. or a matter of convenience to be settled by analogy. under which head or by what language they shall be described! How- ever. most leaves are so constituted as to leave no doubt whether they are simple or compound. I'.U. The leaflets of a compound leaf being homologous with the lobes or segments of a simple leaf, indeed being such segments fully isolated, the two sorts fall under the same tvpes. A pin- nalely veined simple leaf is the homologiie of one kind of com- pound leaf; a radiately veined leaf, of the other. That is, compound leave-, are either i>!ini or ]>i;<>l/ forninon in Ferns. Diyitiiti'-l'liniatr is where the primary division of the petiole is on the palmate or digitate plan ; the secondary, on the pinnate. This seems to he the case in the Sensitive Plant. Mimosa pudica. Fig. 201*. Hut the leaf is here truly Inpinnate with the primary dm.sions very crowded at the apex of petiole. Conjugate- f innate is the same arrangement, with the primary divisions a single pair, at the apex of the petiole, and the leaflets pinnately arranged on these. Digitately or l'ulin«t>-ly Decompound in a nearly regular way is not an uncommon case. Usually, the petiole is succes- sively three-forked, as in Fig. 210, when the leaf is said to l»e bltcnmt? (twice teniate). triternate (thrice ternale). or quadritemate (lour times ter- nate), etc.. according to the nuinher of times it divides, or 2-3— 4:-timestemately compound. The ultimate divisions in such cases of threes are commonly of the pinnately trifoliolate type. 201. Finn* is a convenient name for the partial petioles of a bipinuate leaf, taken together with the leaflets that belong to them. Thus, the Sensitive Plant. Fig. 20'.). has four piniuv. or two pairs; the Honey Locust, Fig. 208, a greater number. When such leaves are still further compounded, the pinna' of higher order, or the ultimate ones, take the diminutive term of PINNTI..K or PINXTLES. The blades these beat- are the LealleN. 202. The Petiole or Leafstalk is a comparatively unessential part of the leaf. It is often wanting (then the blade is .sv.vsVA-) : it may lie absent even in compound leaves of the palmate type, the leaflets rising side by side from the stem. When present, it is usually either round, or half-cylindrical and channelled on the upper side. In the Aspen, it is flattened at right angles with tin- blade, so that the slightest breath of air puts the leaves in motion. Sometimes it is much dilated and nicmbranaeeous at KM!. 210. Quadri-teniivtely compound or ternately decompound leaf of Tbalictruin Cornuti. THEIR STRUCTURE AND FORMS. 100 base, as in many umbelliferous plants ; sometimes it forms a sheath, occasionally it is bordered with appendages, &c. Peti- oles ma}r assume special functions, to be hereafter considered. The woody and vascular tissue runs lengthwise through the petiole, in the form usually of a definite number of parallel threads, to be ramiiied in the blade. The ends of these tlnv:i. as in many Rubiacea-. A notch or fork at the apex often indicates the composition. 206. Sheathing stipules, like those of Polygonum (Fig. 212). are said to be ochreate, or (better) ocreate ; the sheath, thus likened to a leggin or the leg of a boot, is an OCTIREA, as written by Willdeuow, or better OGRE A. 207. The LIGULE of Grasses (Fig. 150) is seemingly a thin and seai-ioiis extension of the lining to the sheath which answers to petiole in such leaves : it projects at the junction of the sheath and blade, there forming a kind of ocrea ; and it is generally regarded as a sort of stipule. 208. Stipels (Stipella) are as it were stipules of leaflets, which are common in certain tribes of Papilionaceous Leguminosne, e. g. in the Phaseolea1. in Wistaria, Locust, &c. ; also in Staphylea. The}7 are small and slender, and, unlike stipules, they are single to each leaflet, except to the terminal one, which has a pair. As leaves furnished with stipules are said to be stij»tfnrkle^ Fig. 215) is obviously the result of a congeni- tal union of ihc bases of the pair by their contiguous edges. Lea\es ,-n, mate in this way by narrow bases are not rare nor remarkable ; but when the two are thus coalescent into one broad ibliareoiis body, giving this appearance of perfoliation, the term connate-perfoliate is used to express it. 2i:'>. Vertical Leaves, those with blades of the ordinary kind, but presenting their edges instead of their faces to the earth and sky, or when erect with one edge directed to the stem and the other away from it. are not uncommon. They prevail in the Australian Myrtacea-. iVc.. and oecur with less constancy in the California!! Manzanitas, and in a great variety of herbs and shrubs. The anomaly involves no exception to the rule that a leaf-blade is always expanded in the horizontal plane, when expanded at all ; for, except in equi- tant leaves, it is the result of a twist of the petiole or of the blade itself.1 In strongly marked cases, or in most of them, the organi/ation of the epidermis and superficial parenchyma and the distribution of the stomata are the same on both faces. 214. Equitnnt Leaves are vertical on a different plan. They are conduplicate, i. e. are folded together lengthwise on their middle, the upper surface thus con- cealed within, the outer alone presented to the air and light. 1 Silpliimn laciniatum. the so-called Compass Plant, and (hardly less so) S. lerchinthinaocnm, arc good instances of the kind, most of the leaves making a half-twist, the radical ones by their long petioles. In the former species, the pimiatelv parted blade occasionally makes a farther twist, so as to bring the upper part into a piano at right angles to the lower. The blades place themselves in various directions as respects the cardinal points ; Init on the prairies the greater number affect a north ami south direction of their edges,— a peculiaritv first pointed out, in the year 1842, by General B. Alvord, I'. S. A. Fir.. 21fi. Kqnitant rivet lr:iv.'s of Iris, with the rootstock. FIG. L'17. A section across lliese leaves at tin- luisc. showing the <'. phvllodia are generally distinguished from true blades by the parallel venation, and alwaj'S by their normally vertical dilatation ; i. e. they, without a twist, present their edges instead of their laces to the earth and sky. The common and most familiar phvllodia are those of Acacias in Australia (Fig. 223, 224), where they form the adult foliage of over 270 out of less than 300 species. The true lamina of these is bipinnate. It appears on seedlings, and occasionally on later growths. Several South American species of Oxalis produce phvllodia. So likewise do our tubular or trumpet- leaved species of Sarraccnia in that portion of the foliage which develops the pitcher imperfectly, or not at all. Indeed, all Sarracenia-leaves are phvllodia with the back in most of them hollowed out into a tube or pitcher; and the terminal hood answers to the blade. § 3. LEAVES SERVING SPECIAL OFFICES. 218. Leaves may serve at the same time both their ordinary and some special use, or even more than one special use. For example, in Nepenthes (Fig. 222) there is a well-developed blade, usually sessile, which serves for foliage, a prolongation of its tip into a tendril, which serves for climbing, then an extraor- dinary dilatation and hollowing of the apex of this into a pitcher fora very special use. and a peculiar development of the apex of this into a lid. closing the orifice during growth. Among the special purposes which leaves subserve, and the study of which connects singularities of morphology with teleology, the most remarkable is that of 2 lit. Leaves specialized for the Utilization of Animal Matter. This occurs in leaves which also assimilate, or do the ordinary work of vegetation ; and the special function is usually taken up by some particular portion of the organ. The details of this LEAVES SERVING SPECIAL OFFICES. Ill subject — which has of late become highly interesting — belong to physiology, and therefore to the following volume, to which 220 all historical references are relegated. Only the morphology of such leaves is here under consideration. 220. As Ascidia or Pitchers, vessels for maceration, &c. These occur in several widely different families of plants. The commonest are those of the Sarracenias, natives of Atlantic North America. They are evidently phyllodia (217), the cavity being a hollowed dorsal por- tion : the wing-like or foliaceous por- tion, ahvays conspicuous and forming the ventral border, makes the whole organ or most of it in the earlier leaves of the tubular species. The pitchers of S. purpurea (Fig. 221, 225), the only species which extends north of Virginia, are open cups, half filled with water, much of which may be rain, in which abundance of insects are usually undergoing macer- 225" ation. In S. variolaris (Fig. 226), the hooded summit, answer- ing to the blade of the leaf, arches over the mouth in such wise FIG. 220. Pitchers of Heliampliora ; 221. of Sarracenia purpurea ; 222. of Nepenthes. 223. A phyllodium of a New Holland Acacia. 224. The same, bearing a reduced corn-- pound blade. FIG. 225. Pitcher-leaves of Sarracenia purpurea ; one of them with the upper parr, cut away. 112 MORPHOLOGY OF LEAVES. as to mostly exclude the rain ; in S. psittaeina (Fig. 227) the inflexed anil inflated hood completely excludes it. The water which these contain is undoubtedly a secretion. All entrap flies, ants, and various insects, which in most species, are lured into the pitcher by a >weetish secretion around or at some part of the orifice.1 Few that have entered ever escape ; most are decomposed at the bottom of the cavity. In Darlingtonia ('alilornica (Fig. 22X), the Cali- fornian representative of Sarraeenia, the inflated hood guards against all access of rain, while the orifice is freely open to flying injects from be- neath ; and a singular two-forked appendage, like to a fish-tail (probably the hoinologue of the blade), overhangs the front. The inner face of this appendage is besmeared with the sweet and viscid secretion which allures insects to the open- ing. In this and in Sarracenia variolaris, the sweet secretion in the early season is continued upon the edge of the wing, forming a saccharine trail which leads from near the ground up to the orifice of the pitcher.2 Fig. 22(> represents pitchers of Heliamphora, a little-known South American representative of Sarracenia. Its wing is narrow and inconspicuous, the mouth widely open and directed upward, and the hood reduced to a minute and upright, probably functionless ap- pendage. In Cephalotus — an anomalous plant of Australia, of uncertain affi- nity -- the leaves for foliage are dilated phyllodia ; among them are others completely transformed into stalked and short pitchers, with thickened rim and a well-lifting lid, hinged by one edge. Fig. 22!). The particular morphology of the parts is not well made out. This sweet secretion, which at times is very obvious in the southern species, has also been detected by Mr. Kdwanl Hiinjess in S. purpurea ; but it is rarely seen, and probably plays no important part in the capture and drowning of the multitude of insects which these pitchers are apt to contain. ' This trail was discovered by Dr. .1. II. .Mellichamp, of South Carolina. See Proe. Am. Association for Advancement of Science, \\iii. 11:] (1871). FIG. 226. Pitcher of Sarracenia variolaris. 227. Same of S. psittaeina LEAVES SERVING SPECIAL OFFICES. 113 221. The pitcher-bearing leaf in Nepenthes has been referred to (218, Fig. 222) : of this there are various species, all of them somewhat woody climbing plants of tropical Asiatic and African islands of the southern hemisphere, some of them familiar in conservatory cultivation. Here the tendril ma}- be regarded as a prolonged extension of the midrib of the blade, and the pitcher, with its hinged lid, as a peculiar development from its apex. The water contained in the pitcher is a secretion, much of which appears before the lid opens ; and a sweetish excretion at the orifice lures insects. The presence of these in the pitcher increases the watery secre- tion in which the animals are drowned ; and this secretion is ascertained to have a certain digestive power.1 222. The aquatic sacs of Utricularia or Bladder- wort are diminutive ascidia, always under water, and with lid opening inward, like a valve, preventing the exit of minute animals entrapped therein.2 Morphologically, they are doubtless leaves or parts of leaves. 223. As Sensitive Fly-traps. The leaves of all species of Drosera or Sundew are beset with stout bristles tipped with a gland, which secretes and when in good condition is covered by a drop of a transparent and very glairy liquid, sufficiently tenacious to hold fast a fly or other small insect. Adjacent bristles, even if not touched, in a short time bend towards those upon which the insect rests, and thus bring their glands also into contact with it. In Drosera filiformis, the leaves are fili- form, with no distinction of petiole and blade. In D. rotundifolia and other common round-leaved species, there is a clear distinc- 1 This was first made out by J. T>. Hooker, and announced in liis address, as President of the British Association for the Advancement of Science, at Edinburgh, 1871. 2 Darwin, Insectivorous Plants, 395. Colin, Beitrage zur Biologic der Pflanzen, 1875. Mrs. Treat, in The Tribune, New York, September, 1874, and Card. Chron. 1875, 303. FIG. 228. Pitcher of Darlingtonia Califoraica. 229. Pitcher of Cepbalotus follicu- laris, -with lid open. 228 114 MORPHOLOGY OF LEA VMS. tion of petiole and blade, and the stalked glands thickly beset the whole upper surface of the latter. A -mall iiix-ct alighting thereon is helpless, and is soon touched by all the glands within reaching distance ; also the blade itself commonly incurves, taking part in the general movement. It lias recently been demonstrated that the captured insect is led upon, and that the plant thereby receives nourishment. Here leaves which do the normal assimilative work of vegetation, but somewhat feebly (having a comparatively small amount of chlorophyll), have also the power and the habit of obtaining ready-organized food by capture, and are benefited by it. L'-JI. Species of Drosera inhabit most parts of the world, and the genus is numerous in species. A near relative. Diona-a. is of a single species, 1). muscipula (Yenus's Fly-trap), inhabiting only a limited district in the sandy eastern border of North Carolina. It is more strikingly sensitive and equally carnivo- rous, but in a different way. It is destitute of stalked and viscid glands. The apparatus for capture and digestion is the two- valved body at the top of each leaf. (Fig. L'.'iO, 231.) If this FIG. 230. A plant of Dion:i-:i inusripula, reduced in size. 231 Three of the leaves, nf almost the nat ural size ; one of them open, the others closed. Probably a fly is never caught by the teeth, in the manner here represented. LEAVES SERVING SPECIAL OFFICES. 115 be taken for the leaf-blade, the part below would be a broadly- winged foliaceous petiole. If the latter be the true blade, the apparatus in question must be reckoned as a peculiar terminal appendage. Both are moderately green, and act as foliage. The specially endowed terminal portion acts also in a decidedly animal-like manner. When either of the three or four slender bristles of the upper surface are touched, the trap suddenly closes, by a movement ordinarily quick enough to enclose and retain a fly or other small insect. The intercrossing of the stout marginal bristles detains the captive, unless it happens to be small enough to escape by the intervening little openings. Otherwise, the sides soon flatten and are brought firmly into contact, and a glairy secretion is poured out from numerous immersed glands : this, with the extracted juices of the macerated insect, is after some time reabsorbed ; the trap, if in a healthy condition, now re-opens and is read}r for another capture. For references to the now copious literature of this whole subject, and for its plrysiological treatment, the succeeding volume should be consulted. 225. Leaves for Storage. Nutritive matter is stored in leaves in many cases, and not rarely in leaves which at the same time are subserving the purpose of foliage. This occurs in all fleshy leaves, to a greater or less extent, according to the degree of thickening or accumulation. The leaves of the Century Plant or Agave, for instance, are green and foliaceously efficient at the surface, while the whole interior is a store-house of farinaceous and other nutritious matter, as much so as is a potato. The leaves of various species of Aloe, Mesembryanthemum, Sedum, and other " succulent" plants (in which a large part of the accumulation is water) are not rarely so obese as to lose or much disguise the foliaceous appearance. Sometimes one portion of a leaf is of normal texture and use, while another is used as a reservoir for the nourishment which the foliaceous part has produced. Fig. 232, a leaf from the bulb of White Lily, the base of which forms one of the bulb-scales, is an instance of the kind.1 The most 1 In Dicentra Cucullaria and (more strikingly from the sparseness of the grains) in D. Canadensis, the matter elaborated in the much dissected blade is conveyed to the very base of the long petiole, and there deposited in a con- FIG. 232. A radical leaf of the White Lily, with its base thickened into a bulb-scale, which is cut across to show its thickness. 116 MORPHOLOGY OF LEAVES. decisive instance of leaves used for storage of food is in that material provision for the nourishment of the embryo in germi- nation, in which the first leaves, the cotyledons. :nv turned to this account. (21-.'57. iVc.) After or \\liile discharging this special duty, the cotyledons may I'ullil their gen- eral oilier, by serving as foliage (as in Maples. Fig. :"!. FIG. 234. A vernal shout of common Barberry, showing a lower leaf in the normal state; the next partially, those still higher completely, transformed into spines. FIG. 235. Solanum jasminoides, climbing by coiling and at length indurating petioles. 118 MORPHOLOGY OF LEAVES. of a pair of stipules into tendrils in Smilax. At least the ten- drils here occupy the position of adnate stipules. The tendrils of Cucurbitaceae are peculiar and ambiguous, on account of their lateral and extra-axillary position and the manner in which the compound ours develop their branches. But they arc doubtless partly if not wholly foliar.1 L'2:i. IVtaloid Leaves, Bracts. Certain leaves, situated near to (lowers, and developing little or no chlorophyll in their paren- chyma, exchange the ordinary green hue and herbaceous texture for the brighter colors and more delicate structure which are commonly seen in and thought to characterize flower-leaves. Such are said to be colored, meaning, as applied to foliage, of some other color than green. As petals are the type of such colored parts, they are said to be petaloid, i. c. petal-like. They are like petals, moreover, in one of the purposes which these sub- serve. (2!M».) Examples of these petaloid leaves are seen in the shrubby .Mexican Euphorbia called Poinsettia, in Salvia splen- clens, most species of Castilleia or Painted Cup, also in the white hood of Calla and Richardia ^Ethiopica (called Calla Lily), and in the four white leaves which subtend the flower-head of Cornus florida, and of the low herbaceous Cornel, C. Canadcn- sis. (Fig. •204.) Such leaves, being in proximity to flowers, and all others which are within a flower-cluster or are borne by flower-stalks, receive the special name of BRACTS. More usually bracts are not petaloid, but different in si/,e or shape from ordi- nary leaves, either by abrupt change or gradual transition. Not uncommonly they are reduced to scales or mere rudiments or vestiges of leaves, of no functional importance. ^.'!0. Flower-Leaves. The morphology of leaves extends not only to "'the leaves of the blossom," more or less accounted as such in common parlance, but also to its peculiar and essential organs, the relation of which to leaves is more recondite. Their morphology needs to be treated separately, and to lie preceded by a study of the arrangement of leaves and of blossoms. 1 Tin- most satisfactory interpretation may In- that of Braun and Wydlcr, adopted by Kichlcr (Bliithendiagranime, i. '504) : that tin- (lower of Ciieur- bita and its peduncle represent tin- axillary branch, the tendril by its side answers to one of the bractlcts (that of the other side being suppressed), and the supernumerary branch springs from the axil of the tendril. This makes of the tendril a simple leaf, of which the branches are the ribs. But the tendril-divisions are evidently developed m spiral order, and in vigorous growths occupy different heights on the tendril-axis. This favors Naudin's view, that the main tendril is cauline, and its divisions leaves. PHYLLOTAXY, OK LEAF-ARRANGEMENT. 119 CHAPTER IV. PHYLLOTAXY, OR LEAF-ARRANGEMENT. SECTION I. THE DISTRIBUTION OF LEAVES ON THE STEM. 231. PHYLLOTAXY (or Phyllotaxis} is the study of the distri- bution of leaves upon the stem and of the laws which govern it. The general conclusion reached is, that leaves are distributed in a manner to economize space and have a good exposure to light, &c., and that this economy on the whole results from the formation of leaves in the bud over the widest intervals between the leaves next below.1 Leaves are arranged in a consider- able varietj" of ways, which all fall under two modes, the Verticillate and the Alter- nate (13), but which ma}T also be termed the Cyclical \ and the Spiral. 232. Alternate leaves are those which stand singly, one after another ; that is, with one leaf to each node or borne 230 on one height of stem. Verticillate leaves are those with two or more at the same height of i stem, circularly encompassing it, i. e. forming' a Verticil or Whorl. Verticillate and whorled are synonymous terms to denote this arrange- ment. These two kinds of leaf-arrangement are commonly ranked as three, viz. alternate, opposite, and whorled. But the opposite is only the simplest case of the whorled, being 1 For the most comprehensive discussion of phyllotaxy in connection with development, and in view of these relations, see Hofmeister, Allgemeine Morphologic, § 11, and Chauncey Wright, Mem. Amer. Academy, ix. 389. FIG. 236, Alternate, 237, Opposite, 238, Verticillate or whorled leaves. 120 PHYLLOTAXY, OR LEAF-ARRANGEMENT. that in which the members arc reduced to two. This case is so much commoner than whorls of three and of higher numbers that it took from the first its special name of opposite, so that in descriptions the phrase "leaves verticillate " implies more than two leaves in the whorl. But it should lie kept in mind that " leaves opposite " is the same as " leaves in whorls of two." '2:>'.'y. The greater number of pluenogamous plants (all but the monocotyledonous class) begin with verticillate leaves, mostly of the simplest kind (z. e. cotyledons opposite): some continue verticillate throughout; some change in the first leaves of the plumule or after the first pair into alternate, and again into verticillate in or toward the blossom, in the interior of which the alternate arrangement may be again resumed. As Nature passes j-adily from the one mode to the other on the same axis, we ma^' expect that the two may be comprised under some common expression. But they have not yet been combined, except by gratuitous or somewhat forced hypotheses; so that for the present they should be treated in morphology as primarily dis- tinct arrangements.1 234. Verticillate or Cyclical Arrangement. Here the leaves occupy a succession of circles, or form whorls around the stem, two, three, four, five, &c., in each whorl. According to the number, the leaves are opposite, ternate, iji«tfcntitfc, (JH!IKII<\ and so on. The characteristic of the individual whorl is that the members stand as far apart from each other as their number renders possible, i. e. they divide the circle equally. Thus, when onjy two, or opposite, their midribs or axes of insertion have an angular divergence (as it is termed) of 180°; when three, of 120° ; when four, U0° ; when five, 72°. 235. The characteristic of the whorls m relation to each other is, that the members of successive whorls stand over or under the intervals of the adjacent ones. In other words, successive whorls ultrnnilc or t/cc/tssafe. This ecoiiomixes space and light, or gives the best distribution which the cyclical system is capa- ble of. And it is in accordance with the general conclusion of Hofmeister's investigation of tin- origin of phyllotaxic arrange- ments in the nascent bud, viz. that new members originate just over the widest intervals between their predecessors next below. Thus, in opposite leaves or whorls of two (Fig. 2.">7), the suc- cessive pairs ili'i'Hssnte or cross at right angles, and so four 1 It is readily seen that whorls may be produced by the non-development of the internotles bet WITH the leaves of a series of two, three, five, or more in alternate order. The ditlieulty is that the members of the next whorl do not follow the rrder that they should upon this supposition. DISTRIBUTION OF LEAVES ON THE STEM. 121 straight equidistant vertical ranks are produced. In ternate or trimerous whorls there are six vertical ranks ; in quaternate or tetramerous whorls, eight vertical ranks, and so on.1 236. The cases in which successive © pairs of leaves do not decussate at right angles, or the members of whorls are not ~ ' exactly superposed to intervals, but as it were wind spirally (as in Dipsacus, many Caryophyllaceae, &c.) , may some of them be explained by torsion of the stem, such as is very manifest in numerous in- stances ; and others may be resolved into 239 instances of alternate leaves simulating or passing into whorls by the non-development of internodes.2 237. Alternate or Spiral Arrangement. Here the leaves are distributed singly at different heights of the stem, and at equal intervals as respects angular divergence. (Fig. 236.) This angu- lar divergence (i.e. the angular distance of an}' two successive leaves) differs in the various kinds of this system of phyllotaxy, but is alwa3*s large enough to place the leaves which immediately 1 These vertical ranks have, by some German botanists, been named Orthostichies ; but this technical Greek is no clearer and no shorter than the equivalent English, which answers every purpose. 2 In Lilium Canadense, superbum, &c., with whorls of variable number of leaves and vague relation to each other (when of the same number some- times the members superposed), and above and below passing into the alter- nate arrangement normal to the family, these whorls are evidently formed of alternate leaves brought together by non-development of internodes. Here may also be mentioned the not uncommon anomaly in Fir-cones, notably those of Norway Spruce, the normal phyllotaxy of which is simply spiral, but in occasional instances the cone is composed of pairs of opposite scales, spirally arranged, /. e. the pairs not decussating at right angles, thus forming double spirals. In the abnormal spruce-cones, the fractions usually observed are •£$ or ^8g, or, as expressed by Braun, (i)^ and (-i)^. Braun's mode of notation for the ordinary succession (i. e. the decussation ) of opposite leaves is (i)i, the | meaning that the two leaves of the pair are half the circumference of the circle apart, the J denoting that each leaf of the succeeding pair diverges one fourth of the circumference from the pre- ceding. Braun finds cases in which pairs (and equally whorls) are super- posed (e.y. certain species of Mesembryanthemum and Euphorbia), these are expressed in this notation by the formula (•£)£, that is, the corresponding leaves of the succeeding pair diverge 180° from their predecessors. He recognizes also some cases of intermediate divergence; such as (£)f in the upper leaves of Mercurialis perennis, (i)^ on certain stems of Linaria vul- garis, (i)^ exceptionally in the leaves of Epilobium angustifolium and the scales of Norway Spruce, (i)-^ exceptionally in the scales of Norway Spruce. See Ordnung des Schuppen an der Tannenzapfen, 376, &c. FIG. 239. Ground-plan diagram of six trirnerous whorls, showing their alternation. 122 PHYLLOTAXY, OB LEAF-ARRANGEMENT. follow each other in the ascending order upon different sides of the axis : it also secures an advantageous spacing of the leaves over the whole length of the axis. Their vertical distance from each other of course depends on the length which the internodo attain, which is a matter of growth and is very variable : but their angular distance is fixed in the kind or numerical plan of the particular phyllotaxy, and is uniform throughout. 238. The leaves are said to be alternate, because they come one after another, now on this side, then on that, as they ascend the stem. The arrangement is said to be spiral, because if a line be drawn or a thread extended from the base or insertion of one leaf to that of the next higher, and so on, taking in all the leaves, it forms a helix, more or less loose or close according to the development of the internodes. (See Fig. 242.) This imagined spiral line ascends continuously, without a break ; and on it the leaves are equably laid down.1 239. Almost ah1 the ordinary instances of spiral phyllotaxy belong to one series, having very simple arithmetical relations. So that this may be taken as the type, and the few others re- garded as exceptions or sometimes as modifications of it. The kinds are simply designated by the number of vertical ranks of leaves: they are technically named by prefixing the proper Greek numeral to the word meaning row or rank. The arrange- ment called Distichous, or Two-ranked, is the simplest and among the com- monest, occurring, as it does, in all Grasses and many other monocotyledonous plants, in Lindens, Elms, and many dico- tyledonous genera. Here the leaves are disposed alternately on '•xactly opposite sides of the stem (as in Fig. 1) : the second leaf being the farthest possible from the first, as is the third from I he second ; the third therefore' over the first, and the fourth over the second, and so on, thus forming two vertical ranks. The angular divergence is here half the circumference, or 1X0°; and the phyllotaxy may be represented by the fraction [.. which desig- nates the angular divergence, while its denominator expresses the number of vertical ranks formed. TristichouS) or Three-ranked, is the next in the series, and is 1 But when we reach a leaf which stands directly over ;\ lower and older one, we say that one set or s/iirc is completed, and that this leaf is the first of a succeeding set or sjiirr. From analogy of such an open spire to the closed cycle of a whorl of leaves, it is not unusual to designate the former likewise :is a o/<-/«. Yet it is better (with Eichler) to restrict that term, and the adjective ryr/W, to vcrticillate phyllotaxy, or to whorls, to which it properly and etymologically belongs. DISTRIBUTION OF LEAVES ON THE STEM. 123 less common, though not rare in monocotyledonous plants. Fig. 240 illustrates it in a Sedge, and 241 is a diagram in horizontal section, as of a bud ; both extending to six leaves or two turns of the spiral. The fraction J- designates this arrangement. The angular divergence, or distance of the axis of the first leaf from the second, and so on, is one third of the circum- ference (or 120°) : conse- quently the fourth leaf comes over the first, the fifth over the second, the sixth over the third, and so on ; that \ is, the leaves fall into three vertical ranks. The spiral character here begins to be manifest, or becomes so by drawing a line on either fig- ure from the axis or midrib of the first leaf to that of the second, and so on to the sixth, forming a helix of two turns.1 Pentastichous, or Five-ranked, sometimes termed the quincuncial arrangement. This is the most common in alternate-leaved dicotyledonous plants. It is shown in Fig. 236 (on a branch of Apple-tree) , and by diagrams, displaying the spiral character, in Fig. 242, 243. The angular distance from the first to the O O second leaf (passing the shorter way) is f of the circumference, or 144°. But the spiral line makes two turns round the stem, on which six leaves are laid down, with angular divergence of f , 1 The line is supposed to follow the nearest way, and the divergence is counted as £, this being the simplest and most convenient. If for any reason the longer way is preferred, then the angular divergence would be expressed by the fraction |. FIG. 240. Piece of a stalk, with the sheathing bases of the leaves, of a Sedge-Grass (Carex crus-corvi), showing the three-ranked arrangement. 241. Diagram of the cross- section of the same. The leaves are numbered in succession. FIG. 242. Diagram of position of six leaves in the five-ranked arrangement : a spiral line is drawn ascending the stem and passing through the successive scars which mark the position of the leaves from 1 to 6. It is made a dotted line where it passes on the opposite side of the stem, and the scars 2 and 5, which fall on that side, are made fainter. 243. A plane horizontal projection of the same; the dotted line passing from the edge of the first leaf to the second, and so on to the fifth leaf, which completes the frura; as the sixth would come directly before, or within, the first. 124 I'lIYLLOTAXY, OR LEAF-AKK AN<; KMKM . 10 and the sixth is the first to come over any one below: {ho seventh comes over the second, the eighth over the third, &c. The leaves are thus brought into five vertical ranks; l,uf these live leaves are laid down on two turns of the helix (the sixth beginning the second revolu- tion) ; the angular divergence of the leaves in order is :, or 144°; the angular distance of the vertical ranks. 72°. This is a very advan- -'•» taij;eons distribution for ordinary foliage on erect or ascending branches. Its formula is f , expressing the angular divergence, the denominator also indicating the; number of vertical ranks, the numer- ator indicating the number of revolutions made in add- ing one leaf to each rank. Fig. 244 illustrates this ar- rangement on a cone of American Larch, the scales of which are homologous with leaves, the numbers in sight are affixed, and those of the whole cone displayed on a plane at the side. Octostichous, or Enj/it-ranL-i',!. a less common arrangement, occurs in the Holly, Aconite, the radical leaves of Plantago. It has the annular divergence of 135°, or f of the cir- cumference. and the leaves in eight ranks, the ninth over the first and at the completion of the third revolution : it is therefore repre- sented by the fraction §. 240. The obvious relations of the fractions ,': • 5!' s- representing the primary forms of spiral phyllotaxy, are that the sum of any two numerators is the numerator of the next suc- ceeding fraction, and the same is true of the denominators ; also the numerator is the same as the denominator of the next but one pre- eecding traction. Following these indications, jj!. «&<•• Now these i •_• ' the series may be extended to /T, FIG. 244. A enne nf Hi,. small-fruited American Lurch (Larix Americana), with the BCalea uuml.ered, rxliil.itin- the live-ranked arrangement. l-'li;. L'lr,. AH otisrt <>r tlie Houseleek, exhibiting the 6-1 3 arrangement; the leaves in sight, numbered, the 1 4th over the- tirst, the lilth over the lith. .Ve FIG. 246. Cotieof Whit,- Pine < I'iiius Strobus) with scales numbered from bottom, and some secondary spirals marked. DISTRIBUTION OF LEAVES ON THE STE.M. 125 cases actually occur, and ordinarily only these.1 The -^ and v/i are not uncommon in foliage. The rosettes of the House- feek exhibit the -f3 or thirteen-ranked arrangement, as also does the cone of Pinus Strobus, the 14th leaf falling over the first. (Fig. 246.) The ^8T is perhaps little less common in foliage upon very short internodes, as likewise are higher ranked numbers ; and in many pine-cones and similar structures ^J and f £ phyllotaxy may be readily made out. This actual series, i, J-, §, f, &c., answers to and may be expressed by the con- tinued fraction, \ + ± T 1 When other instances are detected, they are found to belong to other series, following the same law, such as the rare one of \, \, f , ^. 2 " The ultimate values of these continued fractions extended infinitely are complements of each other, as their successive approximations are, and are in effect the same fraction, namely, the irrational or incommeasurate inter- val which is supposed to be the perfect form of the spiral arrangement. This does, in fact, possess in a higher degree than any rational fraction the property common to those which have been observed in nature ; though practically, or so far as observation can go, this higher degree is a mere refinement of theory. For, as we shall find, the typical irrational inter- val differs from that of the fraction f by almost exactly ToW, a quantity much less than can be observed in the actual angles of leaf-arrangements." "On this peculiar arithmetical property .... depends the geometrical one, of the spiral arrangement, which it represents ; namely, that such an arrange- ment would effect the most thorough and rapid distribution of the leaves around the stem, each new or higher leaf falling over the angular space be- tween the two older ones which are nearest in direction, so as to subdivide it in the same ratio in which the first two, or any two successive ones, divide the circumference. But, according to such an arrangement, no leaf would ever fall exactly over any other ; and, as I have said, we have no evidence, and could have none, that this arrangement actually exists in nature. To realize simply and purely the property of the most thorough distribution, the most complete exposure of light and air around the stem, and the most ample elbow-room, or space for expansion in the bud, is to realize a property that exists separately only in abstraction, like a line without breadth. Neverthe- less, practically, and so far as observation can go, we find that the fractions •f and -f%, -$i, &c., which are all indistinguishable as measured values in the plant, do actually realize this property with all needful accuracy. Thus, | = 0.375, ^ = 0.385, and /T = 0.381, and differ from k [the ultimate value to which the fractions of this series approximate, or what is supposed to be the type-form of them] by —0.007, +0.003, and —0.001 respectively ; or they all differ by inappreciable values from the quantity which might therefore be made to stand for all of them. But, in putting k for all the values of the series after the first three, it should be with the iinderstanding that it is not so employed in its capacity as the grand type, or source of the distributive character which they have, — in its capacity as an irrational fraction, — but simply as being indistinguishable practically from those rational ones." Chaucey Wright, in Mem. Amer. Acad. ix. 387-390. 126 PHYLLOTAXY OR LEAF-ARRANGEMENT, 241 The successive grades of angular divergence of alternate leaves, as expressed in degrees, are £ = 180° § = 144° ^ = 138° 27' 41.54" £ = 120° | = 135° 28j- = 137° 8' 34.29" and so on ; and beyond, if not in the latter cases, the differences become quite too small for determination by inspection. The}" all fall within the £ and £ as to amount of divergence ; and they form a series converging to a deduced t3"pical angle of 137° 30' 28", which, being irrational to the circumference, would place no leaf exactl}" over any preceding one, but alternately and more and more slightly on one and the other side of the vertical, and so on, in an endless spiral. That is, according to Bravais, the ranks in the higher grades tend to become curwiserial, or actually become so; while in the lower grades they are obviousl}' rectiserial. Unless, indeed, there is some torsion of the axis, 03- which the vertical ranks are rendered oblique, as is often the case in cones of the Norwa}" Spruce. But, apart from this, the difference between rectiserial in a high order and curviserial soon becomes inappreciable. Any and all of the higher grades, and practically one as low as the f , secures the utilit3- of the theoretical angle, viz., that " b3" which the leaves would be dis- tributed most thoroughly and rapidly around the stem, exposed most completely to light and air, and provided with the greatest freedom for S3*mmetrical expansion, together with a compact arrangement in the bud." Even in the simpler grades of com- monest occurrence, each leaf (according to Wright) is so placed over the space between older leaves nearest in direction to it as always to fall near the middle of the space, until the circuit is completed, when the new leaf is placed over an old one.1 242. It is to be noted that the distichous or i variety gives the maximum divergence, viz. 180°, and that the tristichous or ^ gives the least, or 120° ; that of the pentastichous or f is nearly the mean between the first two; that of tin- *. nearly the mean between the two preceding, &c. The disadvantage of the two- ranked arrangement is that the leaves are soon superposed and so overshadow each other. This is commonly obviated by the length of the internodes, which is apt to be much greater in this than in the more complex arrangements, then- Co re placing them vertically farther apart ; or else, as in Kims, lieeches, and the 1 This corresponds with Ilofmeister's general rule, that "new lateral members have their origin above the widest iraps between the insertions of the nearest older members." Yet the fact that the character of the leaf- arrangement is laid down at the beginning in the bud does not go far in the way of the mechanical explanation which he invokes. DISTRIBUTION OF LEAVES ON THE STEM. 127 like, the branchlets take a horizontal position and the peti- oles a quarter twist, which gives full exposure of the upper face of all the leaves to the light. The * and f , with dimin- ished divergence, increase the number of ranks ; the § and aU beyond, with mean divergence of successive leaves, effect a more thorough distribution, but with less and less angular distance between the vertical ranks. •24~2a. The helix or primitive spiral upon which the leaves successively originate ascends, sometimes from left to right, sometimes from right to left,1 commonly without change on the same axis, and prevailingly uniform in the same species ; but occasionally both directions occur in the same individual. The earliest leaves of a stem or branch, or the last, are often on a different order from the rest ; or (as alread}- stated) the spiral ma}' change into the cyclical, or vice versa. 243. The relation of the phyllotaxy of a branch to the leaf from the axil of which the branch springs is somewhat various. But in Dicotyledons, the first leaf or the first pair of the branch is mostly transverse ; that is, the first leaves of the branch stand to the right or left of the subtending leaf. In Monocotyledons, the first branch-leaf is usually parallel to and facing the subtending leaf, as shown in Fig. 304. 244. When the internodes are considerably lengthened, the normal superposition of leaves is not rarely obscured by torsion of the axis : indeed, this may equally occur in short internodes, sometimes irregularl}- or in opposite directions, sometimes uni- formly in one. Thus, in Pandanus utilis, or Screw-Pine, of tristichous arrangement, the three compact vertical ranks be- come strongly spiral by a continuous torsion of the axis. The later leaves of Baptisia perfoliata, which are normally distichous, become one-ranked by an alternate twist, right and left, of the successive internodes. 245. When the internodes are short, so that the leaves approx- imate or overlap, it is difficult or impossible to trace the suc- cession of the leaves on the primitive spiral, but it is easy to see which are superposed. The particular phyllotaxy may then be determined by counting the vertical ranks, which gives the denominator of the fraction. But in compact arrangements these vertical ranks are commonly less manifest than certain oblique ranks, w'hich are seen to wind round the axis in oppo- site directions. (See Fig. 245, 246.) These are termed second- ary spirals, also by some parastichies. These oblique spiral 1 That is, of the observer and as Been from without. See p. 51, foot-note. 128 PHYLLOTAXY, OR LEAF-ARRANGEMENT. ninks arc a necessary consequence of the regular ascending arrangement of parts with equal intervals over the circumference of the axis; and, if the leaves are numbered consecutively, their numbers will necessarily stand in arithmetical progression on the oblique ranks, and have certain obvious relations with the pri- mary spiral which originates them, as will be seen by projecting them on a vertical plane. I'l.V. Take, lor example, the 1 :iiT:ingenient, where, as in the diagram annexed to Fig. 211. the primitive spiral, written on a plane sin-lace, appears in the i lumbers, 1. '2. .">. 1. ft. I'., and so on : the vertical ranks thus formed are necessarily the numbers 1-6-11: t-9-14; 2-7-12; 5-10-15: and 3-8-13. I'.ut two parallel oblique ranks are equally apparent, viz. 1-.'!-."). which, if \\e coil the diagram, will be continued into 7-9-1 1-1:5-15 : and also the 2-4-U— S-H) continues into 12-1 I. and so on. it' the axis be prolonged. Here the circumference is occupied by two secon- dary left-hand series, and we notice that the common difference in the sequence of numbers is twyo ; that is, the number of the parallel secondary spirals is the same as the common difference of the numbers on the leaves that compose them. Again, there are other parallel secondary spiral ranks, three in number, which ascend to the right ; viz. 1-4-7, continued into 10-K5 ; 3-6-9-12, continued into 15; and 5-8-11-14, &c. ; where again the common ditl'ercnce, 3, accords with the number of such ranks. This fixed relation enables us to lay down the proper numbers on the leave-, when they are too crowded for directly following their succes- sion, and thus to ascertain the order of the primary spiral series by noticing what numbers come to be superposed in the verti- cal ranks. Thus, in the small cone of the American Larch (Fig. -'14). which usually completes only three heights of leaves, the lowest, highest, and a middle one make a vertical row which faces the observer. Marking this first scale 1 . and count- ing the parallel secondary spirals that wind to the left, we find that two occupy the whole circumference. From 1. we number on the scales of that spiral .'1-5-7, and so on. adding the com- mon difference 2. at each step. Again, counting from the base the right-hand secondary spirals, we find three of them, and therefore proceed to number the lowest one by adding this com- mon difference, viz. 1-4-7-10; then, passing to the next, on which the No. .". has already been fixed, we carry on that se- quence, (5-9, &c. ; and on the third, where No. 5 is already fixed, we continue the numbering, .s-11. ifcc. This gives us in the vertical rank to which No. 1 belongs the sequence 1-6-11, showing that the phyllotaxy is of the five-ranked, or % order. DISTKIBDTION OF LEAVES ON THE STEM. 129 It is further noticeable that the smaller number of parallel sec- ondary spirals. 2, agrees with the numerator of the fraction in this the ?; arrangement; and that this number, added to that of the parallel secondary spirals which wind in the opposite direction, viz. 3, gives the denominator of the fraction. This holds good throughout ; so that we have only to count the number of par- allel secondary spirals in the two directions, and assume the smaller number as the numerator, and the sum of this and the larger number as the denominator, of the fraction which ex- presses the angular divergence sought. For this, we must, how- ever, take the order of secondary spirals nearest the vertical rank in each direction, when there arc more than two, as in all the higher forms. But, in all, it is necessary to count only the most manifest secondary spiral of each direction in order to lay down the proper number on the leaves or scales, and so deter- mine the phyllotaxy.1 In a rosette of the leaves of Houseleek (Fig. 245) and a cone of Finns Strobus (Fig. 24G), the num- bers which can be seen at one view are appended, and in the latter the conspicuous secondary spirals are indicated : one to left with a common difference of 5 ; and two to the right, of which the most depressed and prominent has the common dif- ference of 3, the other, nearest the vertical, the common differ- ence 8. The 14th leaf is superposed to the first, indicating the ^ arrangement. The same conclusion is derived from the num- ber of the higher spirals, the smaller 5 for the numerator, and this added to 8 for the denominator. The mathematical discus- sion of these relations, and of the whole subject of phyllotax}', leads into interesting fields. But this sketch may suffice for botanical uses. 246. Relations of AVhorls to Spirals. Verticillate and alternate phyllotaxy, or whorls and spirals, in all complete exemplifica- tions, are to be considered morphologically as distinct modes, not to be practically homologized into one. Nevertheless, transi- tions between the two, and abrupt changes from one to the other on the same axis are not uncommon, the former especially in the foliage, the latter in the blossom. If the spiral be assumed as the fundamental order, it is not difficult to form a clear con- ception as to how such changes come to pass. A single whorl • 1 In applying this method to the determination of the phyllotaxy of a cone, or any such assemblage of leaves, the student should be warned that, although the cones of Pines and Firs are all normally on the alternate plan (while those of Cypresses are on the verticillate), yet in individual cases (common in Norway Spruce) the cone is plainly made up of pairs of oppo- site scales which are spirally arranged. See note under 236. 130 PHYLLOTAXY, OR LEAF-ARRANGEMENT. may most naturally lie "produced by the non-development of the intcrnodi's between any two, three, or more alternate leaves. Two proximate distichously alternate leaves would thus form a pair : the three leaves belonging to one turn of the spiral in the tristiehous ( \ ) arrangement would compose a trimerous whorl ; the five leaves of the two turns in the pentastichous (V) arrange- ment, a ."(-merous whorl, &c. Verifications of this conception, by whorls breaking up or reverting to spirals, are occasionally met with, and the successive overlapping in spiral order of the members of a trimerous or pentamerous whorl is very common. The few instances among phsenogamous plants in which the leaves are opposite and all in the same plane J (that is, the suc- cessive pairs superposed) ma}' be deduced from the distichous alternate mode becoming opposite without further change, by the simple suppression of alternate internodes. The frequent disjunction of the members of the pair in similar and analogous cases goes to confirm this view. But the characteristic of whorls ordinarily is that proximate whorls alternate, that pairs de- cussate. We cannot homologize this Avith spiral phyllotaxy ; tor in this lies the fundamental difference between the two plans. We can explain it only by a reference to Hofmeister's law, which generally governs leaf-origination as to position, namely, that succeeding leaves appear directly above the intervals between the nearest preceding (241, note) : this gives decussation or alternation of successive pairs or whorls.2 247. Hypothesis of the origin of both. Instead of regarding the spiral path on the stem which connects successive alternate leaves as a purely formal representation, it may be conceived to be the line along which the members in some original form were physically connected, in the manner of a leaf-like expansion 1 As in Loranthus Europicus, &c., according to Braun. See 236, note. 2 This renders the verticillatc an advantageous arrangement, perhaps no less so than the distribution which spiral phyllotaxy effects. Both must be Considered to have been determined by and for tlirir respective Utilities, and to have been independent determinations. For "there is no continuity or principle of eoimeetion between spiral arrangements and whorls " (C'hauncey Wright) ; since, although individual whorls are easily reducible to spirals, each succession is an absolute break of that system. As whorls of four members often (as especially in calyx, bracts, &c.) may and sometimes should be viewed as two approximate pairs, so even the spiral of live members, as in a qiiincuncial calyx, has been conceived to consist of two whorls, one of two, the other of three leaves, the second alternating with the first as nearly as possible. But this appears far-fetched and of loose application. It is much clearer as well as simpler to regard the alternate as the fundamental phyllotaxy, and to deduce individual whorls from spirals, if need be, rather than to imagine spirals as somehow evolved from whorls. DISTRIBUTION OF LEAVES ON THE STEM. 131 resembling a spiral stairway. Upon this supposition, the leaves would be the relics, or rather the advantageous results, of the segmentation of such a frond-like expansion, the segments separated through the development of the stem in length and firmness, and modified in the various adaptations to the conditions of higher vege- table life ; even as leaves themselves are modified into tendrils, bud- scales, petals, or other usefully specialized structures. The type on this conception would be a frond, consisting of an elongating axis with a continuous leaf-blade on one side, and this taking a spirally twisted form. But the frond of Fucaceous Algae, Hepaticae, and the like, is two-bladed. While a one-bladed frond, or with one blade suppressed, might be the original of alternate-leaved spirals, the two-bladed frond, simi- larly broken up, would give rise to the opposite or other varieties of verticillate arrangement.1 248. Fascicled Leaves need to be mentioned here, in order that they may be excluded from phyllotaxy. They are simply a cluster or tuft of leaves, belonging to more than one node, and left in a crowded con- dition because the internodes do not lengthen. They ma}' belong either to the alternate or the verticillate series. In Barberry and in the Larch (Fig. 247), they are evidently alternate ; and they may be inferred to be so in Pines (Fig. 248), or even may be seen to be so in the bud-scales which form the sheath sur- rounding the base of the 2, 3, or 5 foliage-leaves. In Junipers, the leaves of the fascicles are in the verticillate order. 1 This is the conception of the late Chauncey Wright. See his elaborate and most suggestive essay in Mem. Amer. Acad. Arts and Sciences, ser. 2, ix. 379, mainly reprinted in Philosophical Discussions (posthumous), 296-328, in which the whole subject of phyllotaxy is acutely discussed, especially in its relation to questions of origin and developed utilities. His conception FIG. 247. Piece of a branch of the Larch, with two fascicles of leaves, i. e. two very short and stout branchlets, bearing scars of former leaves or bud-scales below, and a dense cluster of leaves of the season at summit. The main axis bears scars from which the alternate leaves of the developed axis of the preceding year separated. FIG. 248. Piece of a branch of Pitch Pine, with three leaves in a fascicle or bundle in the axil of a thin scale (a) which answers to a leaf of the main axis. The bundle is surrounded at the base by a short sheath, formed of the delicate scales of the axillary bud, of which the three leaves are the developed foliage. PHYLLOTAXY, OR LEAF-ARRANGEMENT. SECTION II. DISPOSITION OF LEAVES IN THE BUD. 249. Vernation and .Estivation are terms in general use, under which the disposition ol' leaves in the hud is treated. The first relates to ordinary leaves in this early condition ; the second, to the parts of a flower-bud ; not, however, as respects insertion, or position on the axis, which- is phyllotaxy (231). hut as to 249 -.»> 251 the wa}-s in which they ^— » f\ are coiled, folded, over- ^^r l\ ' JL ^^"*^\ laPPcd, &e., either perse ^Jf I/)/) fvf Cv^yJ OY inter se. Prcefoliation "t^l^i ^^^^S an(l Pr&floration are etymologic-ally better terms, substituted by Richard.1 250. The descriptive J 253 terms which relate to individual leaves or parts, whether of foli- age or blossom, mostly range themselves under the heads of plications or of enrolling, and are such as the following, the sectional diagrams of which are copied from the original figures would make the two plans equally primordial. But the freedom with which these actually interchange on the same axis greatly favors the less hypothetical view that whorls may be condensed spirals. This assumes only the well-known fact that internodes may be completely non-developed. 1 Better formed and more expressive terms: but the Linna\-in ones are most in use, and, though fanciful, are not misleading. In English description, it is as convenient and equally terse to say that the parts arc imbricate, val- vatc, &c., " in the bud." Linnanis, in the Philosophia Botanica, described these dispositions of leaves in the bud under the term i'nllntin, — not a happy name, — but did not treat of them in the flower-bud. Later, in Termini Botanici (Amoen. Ac-ad, vi. 17(52, reprinted by Giscke in 1781), lie intro- duced the words V and ^Kxtinititi in their now current botanical sense, to designate, not the time of leafing and of flowering (spring and summer condition), but the disposition of the parts in the leaf-bud and llower-bud (at least of the petals) as respects foldings, coiling, &c., of single parts, and modes of overlapping or otherwise of contiguous parts. The terminology as regards single leaves, Linna-us fixed nearly as it now remains. That of lea ves or their homologues in connection, and as respects the flower-bud, was very imperfectly developed until its importance (and much of its termi- nology) was indicated by Robert Brown, in his memoir on Proteaceae, 1809, in the Prodromus a year later, and in other publications. /'/y.n'.s (the Greek name) is coming into use as a general term for the folding, &c., of single parts. FIG 'Jl!i-'JM. Linnaean iliagrams of sections of leaves in the bud. 249. Comluph- cate. 250. Plicate or plaited 251. Convolute. 252. Revolute. 253. Involute. 254. Uircinate or Circinal VERNATION, OR PR^EFOLIATION. 133 in the Philosopliia Botanica of Linmeus. They were applied only to foliage, but the}' are equally applicable to floral parts. Leaves, ami all homologous or similar organs, if not simply plane, will be either bent or folded or else more or less rolled up in the bud. The first three of the following terms relate to the former, the remaining terms to the latter. They are as to the mode of packing Plicate or Plaited (Fig. 250) , when folded on the several ribs, in the manner of a closed fan, as in Maple and Currant. This occurs only in certain palmately veined or nerved leaves. Condtipllcate (Fig. 249), when folded lengthwise, or doubled up flat on the midrib, as in Magnolia ; a very common mode. The upper face of the leaf is always within. Reclinate or Inflexed, when the upper part is bent on the lower, or the blade on the petiole, as in the Tulip-tree (the blade of which is also conduplicate) . Convolute (Fig. 251), when rolled up from one margin, i. e. one margin within the coil, the other without, as in Apricot and Cherry. Involute (Fig. 253), both margins rolled toward the midrib on the upper face, as the leaves of Water Lily, Violet, &c. ; also the petals of Steironema and Tremandra. Revolute (Fig. 252), similarly rolled backward from both margins, as the leaves of Azalea and Rosemary. Circinal or Circinate (Fig. 254) , when coiled from the apex downward, as the leaves of Drosera and the fronds of all the true Ferns. Corrugate or Crumpled, as the petals of a Poppy, applies to the irregular crumpling of the otherwise plane corolla-leaves. This is a consequence of rapid growth in length and breadth in a confined space. 251. The Ptyxis (or folding, &c.) of an individual leaf, of which the foregoing modifications are the principal, should be distinguished from the arrangement in the bud of the leaves of a circle or spiral in respect to each other. The interest of the latter centres in the flower-bud, i.e. in aestivation. To this the following exposition is devoted, although sometimes applicable to leaf-buds also.1 252. The disposition of parts in aestivation, in respect to each other, is the result partly of their relative insertion, that is 1 In the succeeding paragraphs, it becomes necessary to presuppose so much knowledge of the flower as is implied in the free use of such terms as calyx and corolla, sepals or calyx-leaves, and petals or corolla-leaves. See, if need be, Chapter VI. Sect. I. 134 I'llYLLOTAXY, OR LEAF-ARRANGEMENT. their phyllotaxy. ami partly of tlu- \va\ in which they comport when their margins meet in growth. Those leave- which are within, or of higher insertion on the axis, will almost necessarily IK- enclosed or overlapped: those which are members strictly of the same whorl or cycle may fail to conn- into contact, or may meet without overlapping at the contiguous margins or apex; yet they may lie overlapped, since they may have grown unequally or some a little earlier than their fellows. Conse- quently, no perfectly clear line can be drawn in the flower between cycles and spirals except by their mode of succession. More- over, {estivation strictly so called should be concerned only with the disposition among themselves of the several members of one whorl, or of one complete spiral. So the alternation of contiguous whorls, as of the three inner with the three outer flower-leaves of a Lily or a Tulip (the alternative activation of DeCandolle) , is a matter of phyllotaxy, not of ;estivatic »n. The latter is properly concerned only with the relations of each three leaves to each other.1 •_'.->:$. The proper aestivations may be classified into those in which the parts do not overlap, and those in which they do. Of the first, there are two kinds, the open (5), the normal imbrica- tion is decussate, two exterior and two interior. This is some- 1 All tin- examples referred to result from alternate or spiral phyllotaxy, the former nf higher series, tin- latter of the £ (Fig. -'•">*. :.'•">!>). and of the'? (Fig. li(><») order. Instead of separating (with I>r< 'aiidolle and others) the g arrangement as different in kind from the imbricate (under the name of quincuncial aestivation), we should count it as a typical case. Otherwise the i arrangement might equally claim a generic distinction, also the f, &c. AESTIVATION, OR PR^EFLORATION. times a clear case of binary instead of quaternary, i. e. to be counted as two pairs of opposite leaves ; yet it may be a single whorl of four, notwithstanding the imbrication. Or these four leaves may even, in some cases, be regarded as a portion of a depressed spiral, say of the f order with one piece omitted, and the others adjusted so as to fill the space. 257. There are various deviations from normally imbricative aastivatiou, especially where the members are five, occurring some in regular but more in irregular flowers, which need not be referred to here. One, for which no specific name is requisite, is a case merely of excessive overlapping in the regular way ; namely, where each piece completely and concentrically encloses the next interior, as shown in Fig. 259, representing three petals of Magnolia Umbrella. This the French botanists have called convolute aestivation, because the individual leaves are involute in a manner approaching the convolute vernation of Linnaeus. Another is the Vexillar, as in the Pea tribe (Fig. 306), where members which should be external have somehow developed as internal, both in calyx and corolla. A third (which has received the usually quite meaningless name of CoMear, spoon-like, and is also that to which most French botanists singularly re- strict the name of imbricative) is a state exactly intermediate between the quincuncially imbricate and the convolute or 261 contorted. In it, one leaf is wholly outside, one wholly inside, and three with one margin inside and the other outside. It occurs under two modifications, viz. with the innermost leaf remote from the outmost (Fig. 261), and with it next to the outermost as in Fig. 262. In view of the intermediate character, we had applied to this the somewhat awkward name of Convolute-imbri- cated To bring Fig. 261 back to the quincuncially imbricate 1 It would not be amiss, therefore, to name one of these modes, viz. that of Fig. 261, Subimbricate, and the other, Fig. 262, Subcom-olitfe. George FIG. 261. Quineuncial imbricate modified toward convolute by one edge of the second leaf developing inside instead of outside of the adjacent edge of the fourth. FIG. 262. Convolute modified toward imbricate by one leaf having a margin inside instead of outside its neighbor. FIG. 263. Convolute, or convolutive, or contorted (twisted) aestivation, in diagram. In these three diagrams, the dark circle above represents the position of the axis, the flowers being axillary. 138 PHYLLOTAXY, OR LEAF-AKKANt; KM KXT. form, we have- only to reverse a single overlapping on the left- hand side of the figure. To restore Fig. 202 to the convolute, we have only to reverse :i single overlapping at the lower right- liand side. Changes like these, or the reverse, are not rare in several species, particularly in the corolla. The normal form and the deviation often occur in dill'erent flowers on the same individual, thus indicating an easy passage between the imbricate aestivation in the proper sense and the 2 .Vs. Conroliite, otherwise called Obvolute or Contorted, or Tiuisled, Fig. 263, and inner circle of Fig. 260. Here each leaf successively overlaps a preceding and is overlapped by a folio wini_r one. all having a slight and equal obliquity of position, so that all alike have an exterior and an interior (or a covering and a covered) margin, and all appear to be as it were rolled up to- gether. This is strikingly so when the parts art- broad and much overlapped, as in Fig. 2G4. Brown included this among the forms of imbricate activa- tion, and so does Eichler, particularly distinguishing it, however, under the name here preferred. The 2&j occasional transitions would justify such classifica- tion. But in most cases it is so uniform, and in the corolla so completely characteristic of whole families (such as Malvaceae, Onagracea-, Apocynaceae, Gentianacea-. I'olenioniacea', &c.), and is so distinct in its nature, that it may well take rank among the primary kinds of aestivation. As to its nature, it is evident that while the imbricate mode (at least the ternary, quinary. &c.) indicates or imitates spiral phyllotaxy (some members be- ing within or with higher insertion than othcis ) . the convolute and the valvate (having all the members of the series on the same plane) answer to verticillatc phyllotaxy, or to whorls instead of depressed spirals.1 The name which this mode of Henslow, in Trans. Linn. Soc. ser. 2, i. 178, proposes to call the former liall imbrir.-itc ; tin- latter (following the faulty French example) is his imbricate proper. The subimbricate mode has two varieties, distinguished by Eichler (in Bliithendiagramme) as •,•, /,>•/><. \vlu-n the lower <>r anterior (/. ,. the pieces next the suhteiidinLT bract or leaf) are successively exterior (as in Fig. L'lil ), ami descensive, when the covering is from the upper side, /. e. from the side next the axis. 1 Still, as those members of a quincunx which normally should be wholly external do -ometimes become internal during their development in the bud, similar changes may be conceived to change a quincuncial into a convolute- disposition; but, to effect this, three out of the live overlapping would have to be reversed. FIG. 264. Convolute (alao called contorted) aestivation of a corolla. AESTIVATION, OR PR^EFLORATION. 139 aestivation ought to bear is not yet well settled, but that of con- volute, here preferred, will probably prevail.1 259. In recapitulation, these principal forms of the aestivation of floral circles may be classified in a synopsis. The}' are : I. those not closed = open or indeterminate : II. closed ; and these 1, with the margins not overlapping = valvate ; 2, with margins overlapping ; «, one or more with both margins covered = imbri- cate ; b, all with one margin covered, the other uncovered = convolute. 260. Plicate or Plaited, when applied to the flower-bud as a whole, is in a somewhat different category. The term is here used for the plaiting of a tube or cup, composed of a circle of leaves combined into one body. It is well marked in 265 see the corolla of Convolvulus and of Datura, and in most of the order to which these belong. In Campanula, these plaits are all outwardly sa- lient and straight (Fig. 265) ; in the corolla of most Gentians, the plaits are internal and straight. In Convolvulus and Datura (Fig. 266-268), the narrow plaits overlap one another in a con- volute way, when they are said to be Supervolute. In the common Morning Glory and some other species of Ipomoea, these plaits are besides spirally twisted or 1 See article entitled " ^Estivation and its Terminology," above referred to. The earliest name is Obvolute, given by Linnaeus to the kind of vernation in which two leaves (conduplicate ones in his diagram) are put together so that one half of eacli is exterior, the other interior. That is just the mode in question reduced to a single pair of leaves, as it is in the calyx of a Poppy. Mirbel is the only botanist who has applied the term to aestivation, and to a circle of more than two leaves, and it has never been adopted in botanical descriptions. It has the disadvantage that the prefix ob to botanical terms means obversely or inversely. Contorted (contorta), in English Twisted, is in early and is the commonest use, and it is sometimes expressive. The objection to it is, that contortion or twisting of the flower-bud often conspicuously oc- curs where there is no overlapping of edges (as in many species of Ipomoea) ; that really no twisting accompanies the overlapping in a majority of cases of this aestivation ; and that when there is a twisting it is not rarely in the direction contrary to the overlapping; so that the contortion needs to be FIG. 265. Cross section of the extrorsely plicate or plaited tube of the corolla of a Campanula in the bud. 266. Same of a Convolvulus (Calystegia), the plaits convolute or supervolute. FIG. 267. Upper part of unexpanded corolla of Datura; the plaits convolute or supervolute. 268. Cross section of the same. 140 PHYLLOTAXY, OR LEAF-ARRANGEMENT. contorted in the opposite direction ; that is, the plaits overlap to the right and arc twisted to the left.1 2<> 1. Direction of Overlapping, &c. This is to be noted in the ternary, quinary, or other forms of spirally disposed imbrication, also in convolute and twisted or contorted activation. It may be either to the right (dextrorse} or to the left (si/n'slrorse). The application of this term depends upon the assumed position of the observer, whether outside or inside. AYe always suppose him to stand outside, in front of the object: so when the over- lapping is from right to left of the observer thus placed, as in Fig. 266, it is sinistrorse ; when from his left to right, as in I-'iir. 267, 268, dextrorse.'2 The direction is generally constant. but in many cases only prevalent, in the same plant or the same species, or even the same genus : sometimes it is uniform or nearly so throughout a whole natural order. separately expressed. To describe the aestivation in such cases as dextrorsum contorta et sinistrorsum torta (or in similar English words), when the overlapping is to the right and the twisting to the left, is at least awkward and cumbrous. Convolute is a fitting name, of occasional early application to this aestivation (as by Jussieu to the petals of Malvaviscus), but without definition in this sense; it has for many years been steadily adopted by the proent writer, is employed by Eichler in Germany, and has recently been adopted by G. Henslow and others in Great Britain. It has. however, the disadvantage of having been used by Linnaeus to express the coiling of single leaves, and in a manner not wholly congruous, but still with one edge outside and the other inside. 1 In our phraseology, dextrorsely convolute and sinistrorsely contorted ; in the current phraseology above referred to, dextrorsely contorted or twisted and sinistrorsely twisted! 2 The reasons for adopting this view (in opposition to the authority of Linnaeus and DeCandolle) are given in note on p. 51. ANTHOTAXY, OR INFLORESCENCE. 141 CHAPTER V. ANTHOTAXY, OR INFLORESCENCE. 262. INFLORESCENCE, a term which would literally denote the time of flower-bearing, was applied by Linnaeus to the mode, that is, to the disposition of blossoms on the axis and as respects their arrangement with regard to each other. ANTHOTAXY, a name formed on the analogy of phyllotaxy, and denoting flower- arrangement, is a better term. The subject really belongs to ramification (83, 14-16), and is also concerned with foliation and with phyllotaxy. It is most advantageously treated apart, immediately preceding the study of the blossom itself. 263. In and near the blossom, both axis and foliage very commonly undergo modification, either abrupt or gradual, giving rise in the former to Peduncles and Pedicels, in the latter to Bracts and Bractlets. 264. A Bract (in Latin Bractea) is a leaf belonging to or subtending a flower-cluster, or subtending a flower, and differing from the ordinary leaves in some respect, usually in shape and size, not rarely in texture and color.1 They are commonly, but not alwa}7s, reduced or as if depauperate leaves, of little or no account as foliage, but some- times of use for protection, sometimes rivalling the highly colored flower-leaves for show, more often insignificant or minute and functionless, sometimes obsolete (as in Cruciferse) , or 1 Bracts of the first order are sometimes called floral leaves (Folia floralia) , or at least these are not well distinguished from bracts. But the term floral leaves is descriptively more properly and usually applied to leaves below the bracts or proper origin of the flower-clusters, yet near them, and un- like the proper cauline leaves. It is a vague term, and is in some danger of being confounded (as it never should be) with another vague term, viz. flower-leaves, or the leaf-like organs of the flower itself. FIG. 269. Bract (spatlie) of Indian Turnip, partly cut away below to show the fleshy spike (spadix) of flowers which it surrounds. 142 ANTHOTAXY, OR INFLORESCENCE. fugacious. Each flower is subtended by (grows from the axil of) u bract in Fig. 277-2.SO, &c. A cluster <>!' flowers is sub- tended by a conspicuous and colored bract iu Fig. '2C>'.>. 270. 271 ; by a circle of colored bracts, imitating white petals, in Fig. 2'.»-l. Si-ATiiE is the name given to such an enclosing bract, or to two or more leaves successively enclosing a flower-cluster. 271 INVOLUCRE is the name given to a circle or spiral collection of bracts around a flower-cluster, as in Cornel (Fig. 294, also in Fig. 280 and 286), or around a single flower, as in Ilepatica and Mallow. A compound inflorescence may have both a general and a partial involucre, one tor the general flower-cluster, others subtending the partial clusters. The name of involucre is then reserved lor the general one ; that of INVOLUCEL is applied to the partial, secondary, or ultimate involucres. P>I;A( TI.KTS (Lat. Hnictcola, diminutive of bract) are bracts of ? secondary or ultimate order. For example, in the slender flower-cluster. Fig. "111. 1> is a bract, subtending each individual flower-stalk ; b' is a bract lei, or bract of secondary order, borne on thai part ial llower-stalk itself. The French naturally translate' the Latin />r axis terminated by a flower (Fig. 281-2*4). which answers to a terminal l»iul. If more ilowers appear. so a- to compose a cluster, they spring from the axils, preferably from the highest axils, and are later. The order of evolution is shown in the figures bv the size of the flower-buds or derive of expansion of the blossoms. Fig. 281 best shows why a determinate or delinite in- florescence is sometimes said to be Desce/t '»;/: Fig. 283 show> why it is called < ',',ilr(fi«jul. the central flower first expanding: Fig. 284 exhibits the lateral or circumferential partial clusters later than the central blossom, and their lateral flowers later than their central. 271. Varieties of Indeterminate or Botryose Inflorescence. The names of most of these have been fixed from the time of Linnaeus, hut defined without reference to the order of evolution of the flowers. They are the Raceme, Corymb, and f'//iM. with ilowers raised on pedicels ; the Spike and Jh-ml. with sessile flowers ; also some modifications of these, notably the A/i/f/tf and the X/xtdix. The raceme may be taken as the type. l!»tri/x is equivalent to racemus, &c. ; and, as the type includes diversity of forms to which the name racemose would seem inapplicable, the term bntryosc, (hot r ////'*<•/,,•„ of Eichler) is bot chosen as the general name of it, and is a good counterpart to cymose for the other type. 272. A Raceme (illustrated in Fig. 272. 277. and by diagram in Fig. 278) is a simple flower-cluster, in which the flowers, on their own lateral or axillary pedicels and of somewhat equal length, are arranged along a relatively more or less elongated rhachis or axis of inflorescence. The common Barberry, Cur- rant. Choke-Cherry and Black Cherry, and Lily of the Valley are familiar examples. 27:5. A Corymb (Fig. 275, 279) is a shorter and broader botryose cluster, which differs from a raceme only in the relatively shorter rhachis and longer lower pedicels; the cluster thus lie- coming flat-topped or convex. The centripetal character is thus made apparent. The greater number of the corymbs of Linnaeus and succeeding botanists are cymes, the central flower first ex- panding. And the term cnri/mhosc or corymb-like is still much used in descriptive botany for a ramification which is mainly of the cj-mose type, and where in strictness the term cymose should be employed. 274. An Umbel (Fig. 280), as in Asclepias. &i-., differs from a corymb onby in the extreme abbreviation of the rhachis or axis of inflorescence, and the general equality of the pedicels which thus all appear to originate from the apex of the peduncle, and THE BOTKYOSE TYPE. 147 so resemble the rays of an umbrella ; whence the name, and whence also the pedicels or partial peduncles of an umbel are termed its Rays. The bracts, brought by the non-development of internodes into a depressed spiral or apparent (or sometimes real) whorl, become an involucre. (264.) An umbel or any similar cluster when sessile (without a common peduncle) , and the parts crowded, is sometimes called a Fascicle (or the pedicels said to be fascicled) ; but this term has been differently defined. (280.) It is better not to use it for any special kind of inflores- cence, but simply in the sense of a bundle of whatever sort. This will accord with the sense in which it is applied to an aggregation of leaves. (248.) 275. A Head or Capitulum (Fig. 285) is a globular cluster of sessile flowers, like those of Red Clover, Button-bush, and Plane-tree. The pedicels need not be absolutely wanting, but only very short. An umbel with pedicels much abbrevi- ated thus passes into a head, as in Eryngium, &c. And a head with rhachis elongated passes into a spike. The short rhachis of a head very commonly takes the name of receptacle. (265.) The whole may be subtended by con- spicuous bracts forming an involucre (264) as in Fig. 286, or may be destitute of any, as in Fig. 285. On ac- count of the compactness and mutual pressure under growth, the bracts among the flowers of such heads (normally one subtending each blos- som) are apt to-be rudimentary, reduced to little scales, or abortive, or completely wanting. In the latter case, the recep- tacle is said to be naked (nude} , i. e. naked of bracts : when they are present, it is paleate or chajfy, A peculiar sort of head, not undeserving a special name (though this is not necessary in descriptive botany) , is the ANTHODIUM, the so-called Compound Flower of the earlier botanists, which gives the name to the vast order of Compositae. FIG. 285. Cepbalantbus occidentalis, the Button-bush; a pair of leaves, and a terminal peduncle bearing a dense head (capitulurn) of flowers. 148 ANTHOTAXY, OR INFLORESCENCE. The name means "resembling a flower." Although it has all the characters of a true head, the resemblance to a flower is remarkably striking, the involucre imitating a calyx, and the strap-shaped (ligulate) corollas of the several flowers imitating the petals of a single blossom. In some (such as Dandelion andtheCichory. Fig. I'M',), all the flowers of the liead bear these petal-like corollas; in more (such as Aster, Sun- Mower, and Coreopsis, Fig. 2-S7), only an outer circle of Mowers does so: the remain- der, smaller and filling the centre (or (Jink), may by the casual observer be taken for stamens and pistils, and further the deception. The rhachis or receptacle of a head of this kind is commonly deprived, bearing the Mowers on what then becomes the upper surface, which adds to the imitation.1 SVCONIUM. This name, given to the Fig-fruit, should be here referred to, as it is a sort of inflorescence, of the general nature of a. head, but with receptacle external and flowers enclosed The receptacle of .-in . I ;///„„//,///, has heeii termed < "inuiuthinm or Phor- anthium; ami its involucre, a Periphoranthium or Peridinium. The head has likewise been named a Cephalanthium. FIG. 286. Flowering branch of Cichory, with two heads of ligulate flowers, of natural size. FIG. 287. Vertical section of a head of flowers of a Coreopsis. THE BOTRYOSE TYPE. 149 within. See Fig. 657-659 (683), where its morphology is ex- plained and illustrated. Viewed as an inflorescence, it has also been named a HYPAXTHODIUM. 276. A Spike is a cluster of sessile (or apparently or nearly sessile) lateral flowers on an elongated axis. It may be de- fined by comparison, as a head with the rhachis lengthened (indeed a young head often becomes a spike when older) , and equally as a raceme with the pedicels all much shortened or wanting. A common Mullein and a Plantain (Plantago, Fig. 290) are familiar ex- amples. Two modifications of the spike (or sometimes of the head) gen- erally bear distinct names, although not distinguishable by exact and con- stant characters, viz. : - SPADIX, a spike or head with a fleshy or thickened rhachis. The term is almost restricted to the Arum family and Palms, and to cases in which the inflorescence is accompanied by the peculiar bract or bracts called a spathe (Fig. 269-271). But the two do not always go together : in Acorus and Orontium there is properly no spathe to the spadix ; while in the Iris family the bracts are said to form a spathe, and there is no spadix. In Palms, the principal reason for naming the inflorescence a spadix is its inclusion in a spathe before anthesis. FIG. 288. A slice of Fig. 287, more enlarged, with one tubular perfect flower (a) left standing on the receptacle, and subtended by its bract or chaff (!>); also one ligulate and neutral ray-flower and part of another (c, c) : in rf, d, the bracts or leaves of the involucre are seen in section. FIG. 289. Catkin of White F-irch. 290. Young spike of Plantago major. 150 ANTHOTAXY, OR INFLORESCENCE. AMENT or CATKIN. This is morel}- that kind of spike with scaly bracts borne b}' the Birch (Fig. 289), Poplar, Willow, and, as to one sort of flowers, by the Oak. Walnut, and Hickory. which are accordingly called amentaceous trees. Catkins usually fall off in one piece, after flowering or fruiting. All true catkins are unisexual. '111. Any of these forms of simple inflorescence may be com- pounded. Racemes may themselves lie disposed in racemes, spikes in spikes (as in Triticum) . heads be aggregated in heads, umbels in umbels, coombs may be eorvm- bosely compound, -rayed cymes, &c., may be founded upon alternate leave* with shortened internodes, the rays or peduncles axillary to them thus brought into an apparent whorl. Bravais distinguished cymes as mii/h'/itn-oiix, with three or more lateral axes; biparous, with two; and itni/>itr\ to show why thi> i- not a true raceme. But the other bract of the pair, upon that supposition, is unaccountably empty : the successive angular divergence of each joint of the axis of inflorescence in the younger part, \viiicli commonly runs into a coil, finds explanation in the view that each portion is Hie lateral branch from the axil of the si il (tend- ing leaf: and occasionally the other axil produces a similar one, thus revealing the eymose character. When the bract from the axil of which the missing branch should come disappears also, as sometimes it does, and uniformly on the same side, a state of things like that of the upper part of Fig. 297 occurs. The- same figure may serve for the arrangement corresponding to that of Fig. 206, only with alternate leaves. But then, close as the imi- tation of a raceme here is, the position of each flower in respect to the bract supplies a criterion. While in a true raceme the flower stands in the axil of its bract, here it stands on the oppo- site side of the axis, or at least is quite away from the axil. 282. S.vmpodial forms. The explanation is that the axis of inllorescence in such cases, continuous as it appears to be. is not a simple one, is not a monopode, but a sympode (110, 116, notes), i.e. consists of a series of seemingly superposed inter- nodes which belong to successive generations of axes : each axis bears a pair of leaves (Fig. 296) or a single leaf (Fig. 2!»7). is continued beyond into a peduncle (or pedicel in these instances), and is terminated by a flower. From the axil promptly springs a new axis or branch, vigorous enough soon to throw the adjacent pedicel and flower to one side: this bears its leaf or pair of leaves, and is terminated like its prede- cessor with a flower ; and so on indefinitely. The fact that the alternate leaves or bracts are thrown more or less strict Iv to one side and the1 flowers to the other, in Fig. 297, shows that these leave*, do not belong to one and the same axis; for alternate leaves are never one-ranked or disposed preponderatingly along one side of an axis, as in this diagram, and as is seen in the inflorescence of a Ilouseleek. &c. '2*'.\. A further diflicnlty in the morphology of clusters of this class comes from the early abortion or complete suppression of bracts. This is not unknown in liotryose intloivseeiiee. occurring in the racemes of almost all Crucifenv : it is very common in the t-ymose of all varieties, and especially in the uniparous ones in question, which characterize or abound in l>orraginace;e. Ilvdro- phvllacea'. and other natural orders. In some genera or species, the bracts are present, or at least the lower ones ; in others, THE CYMOSE TYPE. 155 absent ; in some, either occasionally present or wanting in the same species or individual. It is only by analogy, therefore, and by a comparison of allied plants, that the nature of some of these flower-clusters can be made out. With the botanists of a preceding generation, these one-sided clusters were all described as racemes or spikes. Botanists still find it convenient to con- tinue the use of these names for them in botanical descriptions, adding, however, as occasion requires, the qualification that they are false racemes or spikes, or cymose racemes, and the like ; or else, by reversing the phrase, with stricter correctness they call them racemiform or spiciform cymes, &c. 284. Commonly these false racemes or spikes (or botr}roidal cymes, if we so name them) are circinate or inrolled from the apex when young, in the manner of a crosier, straightening as they come into blossom or fruiting. Likening them to a scorpion when coiled, the earlier botanists designated this as scorpioid. As the coil is a helix, it has also been named helicoid.^ The flowers are then thrown, more or less strictly, to the outer side of the coiled rhachis, where there is room for them ; and so these false racemes or spikes are secund or unilateral. The particular anthotaxy and phyllotaxy of the various sympodial and botryoi- dal forms of cymose inflorescence become rather difficult ; and the sorts which have been elaborately classified into species (and have no little morphological interest) are connected by such transitions, and are based on such nice or sometimes theo- retical particulars, that the terminology based on them is seldom conveniently applicable to descriptive botany, at least as to sub- stantive names. 285. One of the latest and simplest classifications of cymes is that of Eichler in his Bliithendiagramme.2 1 Scorpioid and Helicoid have been carefully distinguished by later rnorphologists, on account of some difference in the mode of evolution and arrangement of the flowers along one side of the rhachis, by which they become two-ranked in scorpioid, one-ranked in helicoid. But practically the two kinds of clusters are not always readily discriminated ; and in gen- eral terminology a single name, with subordinate qualifying terms, is suffi- cient. Scorpioid is the older and commoner one, therefore the most proper to be used in the generic sense. 2 CYMOSE TYPE (classified without reference to bracts, which are so often wanting) ; divided into a. Lateral axes three or more: PLEIOCHASIUM, the multiparous cyme oi Bravais. £. Lateral axes two : DICHASIUM, the biparous cyme of Bravais. y. Lateral axis one • MONOCHASIUM, the uniparous cyme of Bravais. The latter, or the corresponding divisions of the preceding sorts, may be divided as follows : 156 ANTHOTAXY, OR INFLORESCENCE. 28G. Sundry complications and obscurities are occasionally encountered in anthotaxy or phyllotaxy. which cannot here be * Lateral axes transverse to the relatively main axis. 1. Lateral axes in successive generations always falling on the same side of the relatively main axis : Sciiu.vri'.Ki. [screwlikc] or BOSTKYX [ringlet or curl], the //////»//-f the Drepanium. 303. Ground plan, the flowers evolved in succession, from left to right. FIG. 304. Diagram showing the position of bractlet or first leaf on a branch in Monocotyledons: a is the primary, a/ the secondary axis; b is bract, and !>' bractlet. THE CYMOSE TYPE. 157 explained, except through full details : such as flowers standing by the side of a leaf, or a small leaf by the side of a larger one, The transverse or oblique position of secondary axes or peduncles, as in Eichler's first two species, brings the flowers of the false raceme or spike out of line of the sympodial axis and bracts, neither in the axils, as in true racemes, nor opposite them, as in the Rhipidium and Drepanium, but on one side of this plane or the other. This is most common in Dicotyledons (in Drosera, Sedum, Sempervivum, and Hyoscyamus, in Borraginacese and Hydrophyllaceas, &c.), and is not rare in Monocotyledons, especially with tristichous phyllotaxy, as in Tradescantia. In the Bostryx, Fig. 299, the bractlet is anterior or falls on the same side as the bract, or, in other terms, the successive bracts are all on one side, the inner side, of the helix ; and the Drepanium (Fig. 301) is like it: this is the helicoid cyme of Bravais, &c., and its flowers are commonly one-ranked. In the Cincinnus or true scorpioid cyme (Fig. 298), and equally in the Rhipidium (Fig. 300), the bracts fall alternately on opposite or different sides of the sympodial rhachis, because the single bract (&') of each successive secondary axis (a') stands next the axis (a) and over against the bract (6) of the generation preceding. The flowers in these generally fall into two parallel ranks (conspicuously so when crowded) on the upper side of the rhachis, on which, in the cincinnus or true scorpioid cluster, they are usually sessile or nearly so (or spicate), as is well seen in Heliotropes, and in very many Borraginaceous and Hydro- phyllaceous species, in Houseleek, Tradescantia erecta, &c. This comes through antidromy, that is, the phyllotaxy of each successive axis of the sympode (with its one bract, or by suppression without it ) changes direction, from right to left and from left to right alternately. Fig. 301 is a plan of this two-ranked unilateral arrangement. When not too crowded, both Cincinnus and Rhipidium are apt to have a zigzag rhachis. These two last-mentioned kinds are so gen- erally alike in character, as are equally the Bostryx and the Drepanium, that the four spe- _ cies may as well be reduced to two. As these v\ V / -^ \ severally include the scorpioid and the helicoid j _ ^ Jt I uniparous cymes of modern anthotaxy, these terms may be retained to designate them. Or, if other terms in use be preferred to scorpioid and helicoid, the form with two-ranked flowers may be denominated Cincinnal, that with single- ranked BostrychoidaL But in neither type is the rhachis always coiled up, although commonly more or less so in the undeveloped state. While these forms generally imitate racemes or spikes, it will be noted that Fig. 300 specially imitates a corymb in form and in seeming acropetal or centripetal evolution. And when, as in this figure, the bracts arc all absent, no obvious external difference remains. FIG. 305°. Ground plan (from Eichler's Bluthendiagramrne, i. 38) of the scorpioid inflorescence of Tradescantia erecta, between bract (B) below and axis (a) above: I., II., III., &c., the successive flowers: i* is the bractlet of the first and bract of the second flower, and so the others in succession up to v5 and a small undeveloped one beyond. The figure 1 affixed to each flower indicates the first floral leaf. 158 ANTHOTAXY, OR INFLORESCENCE. or a pedicel or peduncle above and out of connection with the leaf which should subtend and accompany it.1 287. Mixed Inflorescence is not uncommon. This name is given to clusters or niiiiilic:itions in which the two types are con- joined. Being heteromorphous, they are almost necessarily com- pound, the t\\o types belonging to different orders of ramification. But under it may be included cases of comparatively simple iullorescence, at least in the beginning, some of which nearly fuse the two types into one. In the Teasel ( Dipsacus), an appar- ently simple head or short spike comes first into flower at the middle, from which the flowering proceeds regularly to the base. Had it begun at the top. it world answer to Fig. 281, which, blossoming from above downward by simple uniflorous lateral axes along a monopodial primary axis, is a simple racemiform cj'me. while it may also be called a reversed or determinate raceme. Something of this sort may be seen in certain species of Cam- panula, with virgate inflorescence, the terminal blossom earliest, the others following irregularly, or partly downward and partly upward. In C. rapunculoides, when rather depauperate and /;lie inflorescence simple, the evolution is that of a true raceme, except that a flower at length terminates the axis and develops earlier than the upper half of the raceme. In Liatris spieata and its near relatives, the heads, on the virgate general axis, come into flower in an almost regular descending order, or are reversely spicate. If in Fig. 2M the lower pedicels were prolonged to the level of the upper, a simple coiymbiform cyme would l»e seen, with simple centrifugal evolution, that is, regularly from the centre to the margin ; this is the counterpart of the rhipidium or fan-shaped cyme, of Fig. 300, in which the evolution of the blossoms is as regularly centripetal. The explanation of the paradox is not far to seek. 1 The position of a pedicel at tin- side of a bract in falsi- racemes is ex- plained in the foregoing note. It may occur in true racemose inflorescence by the reduction of sessile secondary racemes down to an umliel of two flowers, transverse to the bract (as in many species of Dcsnmdium), and thus seemingly lateral to it, or to a single (lower on the right or left of it. The coalescence of a pedicel to the axis for a considerable height above the subtending bract in a simple inflorescence, or above the last leaf in a sympodial one (i 'n/n -im/i \r< in-, of Schimper), is common. So likewise bracts or leaves may be for a good distance aduate to sympodial shoots, whether peduncles or leafy tlowerless branches. This (named rccnulescence by Schimpcr) is of most frequent occurrence in Solanacea' (in Datura, Atropa, most species of Solanum, &c.), and is the explanation of their so-called i/i niiinili- leaves, where ;i large leaf (really belonging lower down) has a small leaf by the side of it. See Wydler in Hot. Zeit. ii. (389, &c., Scndtner in Fl. Bras. x. 183, and Eichler, Bluthend. i. 199. THE CYMOSE TYPE. 159 288. Compound mixed inflorescence is very various and com- mon ; but the combinations have hardly called for special terms, being usually disposed of by a separate mention of the general and of the partial anthotaxy, or that of the main axis of inflo- rescence and that of its ensuing ramification.1 In Composite, for instance, the flowers are always in true heads, of centripetal evolution. The heads terminate main stems as well as lateral branches, so that they are centrifugally or cymosely disposed. The reverse occurs in all Labiate and most Scrophulariaceae, where the flowers, when clustered, are in cymes, but these cymes are from axils, and develop in centripetal order. It is this arrangement which mainly characterizes the THYRSUS. A compound inflorescence of more or less elongated shape, with the primary ramification centripetal or botryose, the secondary or the ultimate centrifugal or cymose. To the defini- tion is generally added, that the middle primary branches are longer than the upper and lower, rendering the whole cluster narrower at top and bottom, and sometimes that it is compact = but these particulars belong only to typical examples, such as the inflorescence of Lilac and Horsechestnut. In the former, the thyrsus is usually compound. A loose thyrsus is a MIXED PANICLE. It is seldom that a repeatedly branching inflorescence of the paniculate mode is of one type in all its successive ramifications. Either the primarily centripetal will become centrifugal in the ultimate divisions, or the primarily centrifugal will by suppression soon run into false botryose forms, into apparent racemose or spicate subdivisions. 80 that the name Panicle in terminology is generally applied to all such mixed compound inflorescence, as well as to the homogeneously botryose. (278.) VERTICILLASTER is a name given to a pair of opposite and sessile or somewhat sessile cymes of a thyrsus or thyrsiform inflorescence, which, when full, seem to make a kind of verticil or whorl around the stem, as in very many Labiatse. The name was originally given to each one of the pair of cymes ; but it is better and more commonly used to denote the whole glomerule or false whorl produced by the seeming confluence of the two clusters into one which surrounds the stem. 1 Guillaud (in his memoir on Inflorescence, published in Bull. Soc. Bot. France, iv.) proposes to designate as Ci/mo-Botryes the mixed inflorescence composed of cymes developed in botryose order, i. e. the thyrsus ; and Botry-Cymes, the reverse case of racemes, &c., cymosely aggregated. For the former, the old name thyrsus serves appropriately and well. 160 ANTHOTAXY, OR INFLORESCENCE. 289. The Relations of Bract, Bractlet, and Flower should here be noticed, although the subject in part belongs rather to the section on Floral Symmetry. (31.").) •JIM). Anterior and Posterior, otherwise called Inf<>rinr and Siijicrior. and therefore Lower and f'jyx'r,1 are primary relations of position of an axillary llower with respect to subtending bract and the axis to which the bract pertains. The llower is placed between the two. The portion of the flower which faces the subtending bract is the anterior, likewise called inferior or l.">. the parts are all in the median plane : in Fig. 306, the bractlets, &', b1, are lateral or collateral, or (being in the opposite plane) traiisrerse. 292. Position of Bractlets. The rule has already been laid down (285) that the first leaf of an alternate-leaved secondary axis is in Monoctyledons usually median and posterior, that is. farthest away from the subtending leaf (as in Fig. 301. .">05) ; in Dicotyledons, lateral or transverse. When these secondary axes are one-flowered peduncles or pedicels, the leaf or leaves (if any) they bear are bractlets.'1 Commonly there is only this 1 Not (with propriety, although the terms have been so used) t-.rti ,-inr or nut,',- for the anterior, and inti r/nr or inner for the posterior position. These terms should he reserved for the relative position on the axis of successive circles or parts of circles, spirals, &c. Covering or overlapping parts are exterior or outer in respect to those overlapped. 2 Latin llnniml,! : not that they are small bracts, but bracts of an ulti- mate axis. In axillary inflorescence, the distinction between bractlet and bract is obvious: in ease of a solitary terminal flower, there is no ground of difference: in terminal or cymose inflorescence, the difference is arbitrary ; but we may restrict the term braetlet to the last bract or pair. German botanists mostly distinguish between bracts, as a leaf subtending a flower or cluster, and bractlets, by terming the former a Deckblatt, and the FIG. 306. Diagram (cross section) of papilionaceous flower and its relation to axis (a), bract (b), and l.ructlets (ft', 6'). THE CYMOSE TYPE. 161 posterior one to a simple axis in Monocotyledons, and two transverse ones in Dicotyledons, i. e. one to the right and the other to the left of the subtending bract, Fig. 306, If b' . When the latter form a pair, the}' are perhaps always truly transverse ; when alternate, they stand more or less on the opposite sides and transverse. When more than one in Monocotyledons, the}- may become either median or transverse, or even intermediate. The relation of bracelets or bract, that is, of the last leaves of inflo- rescence, to the first of the blossom, might be considered either under Phyllotaxy or under Floral Symmetry. In general, it may be noted that successive members stand over the widest intervals ; l in other words, that the first leaf of the flower is as far away as may be from the highest bractlet. For instance, when there is a single and posterior bractlet, as is common in Monocotyledons, the first leaf of the flower is anterior, the next two right and left at 120°. When there is a single and lateral bractlet and five leaves in the first circle of the flower (which occurs only in Dicotyledons) , the first leaf of this circle is either exactly on the opposite side from the bract, or at a divergence of two fifths, the latter falling into the continuous spiral. When with a pair of bractlets, right and left, the first flower-leaf is at ^ divergence from one (the uppermost) of them when the circle is of three, or at f when of five members, or near it ; but with many exceptions.2 A tabular view of the kinds of inflorescence and their termi- nology, serving as a key, may aid the student.3 latter, being the leaves which the new axis first bears, Vorblatter, which is also the name they apply to primordial leaves in germination. 1 In accordance with Hofmeister's law ; but (as Eichler remarks) not to be explained on his mechanical principle of production in this place because of the greater room : for the position of the first member of an axillary flower is mostly the same as regards the subtending bract when the bractlets are wanting. '• When bractlets are wanting, the leaves of the first floral circle if two are right and left ; if three, two lateral-posterior and one anterior ; when five, the odd one commonly in the median line, either anterior or posterior. 3 INFLORESCENCE is either PURE, all of one type, or MIXED, of the two types combined. The Types are: I. Main axis not arrested and terminated by a flower. Indeterminate, Indefi- nite, Acropetnl or Ascendinr/, Centripetal, or BOTRYOSE. II. Main and lateral axes arrested and terminated by a flower. Determinate, Definite, Descending, Centrifugal, or CYMOSE. 162 ANTHOTAXV, Oil 1NFLO11ESCENCE. I. BOTRYOSE TYPE. 1. Simple, with lateral axes unbranched and terminated by a single flower, and I lowers on pedicels, ()t -omen hat iM|n;il length on a comparatively elongated axi-. K.M i MI . Tlii- lower oni's longer than the upper, and main axis short, . ('m:YMi:. ()t nearly ec|iial length mi an undeveloped main axis, . . UMI-.II.. Flowers -e-sil,. on a very short main axis, HKAI>. Flowers sessile on a comparatively elongated main axis, . . . SIMM. A fleshy spike or head is a SI-.UMX. A scaly-bracted spike is an AMKNT or CATKIN. 2. Com/n:/ii/'/. with lateral axes branched once or more, bearing clusters instead of single tlowers. Irregularly racemose! v or corvmbosely compound, . . PANICLE. Homogeneously and regularly compound, as Karemrs in a raceme, ( DM mi M. KACKME. Corymbs corymbose ('UMI-IUMI <'<>I:YMR. Umbels in an umbel, COMPOUND UMBEL. Spikes (.picate, COMPOUND SPIKE. Homogeneously compound, the secondary ramification unlike the primary, as Heads racemose, Umbels spiked, Spikes panicled, &c. II. CYMOSE TYPE. 1. Simple, with terminal axis of each generation one-flowered. Mmopodial, the axis of each generation evidently re- solved into branches, TRUE CYME. These more than two, PfetocAottKm or MuLTiPAROUfl Cnot These onlv two, . . . Diclirrsium, l)i< lini«iii,ni* or BITAROUS CYME. Sympodial, the apparently simple axis continued by a succession of new axes standing end to end, Munuchnsiiimj False Raceme or Spike, Botryvse or UNIPAROUS CYME. Flowers one-ranked on one side of rhachis, HKI.K tt . with unbranched one-flowered lateral axes (287), such as ... PARTLY UEVKKSED SPIKES OR RACEMES. 2. Compound, of various combinations, of which there are name's for the subjoined : — Primary inflorescence botryose. with axis elongating; secondary eyinose, THYRSUS. Pair of such opposite cymes seemingly confluent round the main axis, . VEKTICILLASTER. Panicle with some of the ramifications cymose, . . . MIXED PANICLE. THE FLOWER. 163 CHAPTER VI. THE FLOWER. SECTION I. ITS NATURE, PARTS, AND METAMORPHT. 293. FLOWER-BUDS are homologous with (morphologically an- swering to) leaf-buds, and they occupy the same positions. (266.) A FLOWER is a simple axis or a terminal portion of one, in a phffinogamous plant, with its leaves developed in special forms, and subservient to sexual reproduction instead of vegetation. 294. In passing from vegetation to reproduction, it is not always easy to determine exactly where the flower begins. The same axis which bears a flower or floral organs at summit bears vegetative leaves or foliage below. Or when it does not, as when an axillary flower-stalk or pedicel is bractless, the change to actual organs of reproduction is seldom abrupt. Usually there are floral envelopes, within and under the protection of which in the bud the essential organs of the flower are formed. Some or all of these protecting parts, in very many flowers, are either obvious leaves or sufficiently foliaceous to suggest their leafy nature ; and even when the texture is delicate, and other colors take the place of the sober green of vegetation, the}' are still popularly said to be the leaves of the blossom. These pro- tecting and often showy parts, though not themselves directly subservient to reproduction, have always been accounted as parts of the flower.1 Between the lowest or outermost of these and the bractlets and bracts there are various and sometimes complete gradations. The axis itself occasionally undergoes changes in such a way as to render the determination of the actual beginning of the flower somewhat arbitrary. Moreover, the flower itself is extremely various in different plants, in some consisting of a great number of pieces, in others of few or only one ; in some the constituent pieces are separate, in others combined. The flower is best understood, therefore, by taking some particular specimen or class of flowers as a representative 1 Indeed, the colored leaves, or envelopes in whatever form, essentially were the flower in most of the ante-Linnasan definitions (that of Ludwig excepted), as they are still mainly so in popular apprehension. 164 THE FLOWER. or pattern, and especially some one which is both complete and morphologically simple. 295. Such a flower consists of two kinds of organs, viz. the Protecting Organs, leaves of the blossom, or floral envelopes, which, when of two sets, are CALYX and COROLLA ; and the Essential Reproductive Organs, which co-operate in the production of seed, the STAMKV- and PISTILS. 296. Floral Envelopes, Perianth, or Perigone,, the floral leaves or coverings. The former is a proper Knglish designation of these parts, taken collectively. But in descriptive botany, when- a single word is preferable, sometimes the name perianth (Lat. ppn'it/if///H>n), sometimes that of perigone (or perigoniuui) . is used. Peri(i)itJi!inn.} a Linna>an term, has been objected to, because it etymologically denotes something around the flower : but it seems not inappropriate for the envelopes which surround the essential part of the flower. Perigonium, a later term, has the advantage of meaning something around the reproductive organs, which is precisely what it is. Neither name is much used, except where the perianth or perigone is simple or in one set (when it is almost always calyx), or where it is of two circles having the gen- eral appearance of one and needing descrip- tive treatment as such, a- in the petaloidcous Monocotyledons. It is also used where the morphology is ambigu- Generally, the floral envelopes are treated distinctively as calyx and corolla, one or the other of which (mostly the corolla) may be want ing. •_".»7. The Calyx is the outer set of floral envelopes. That is its only definition. Commonly it is more herbaceous or Ibliaceous than the corolla, and more persistent, yet sometimes, as in the Poppy family, it is the more deciduous of the two. Not rarely it 1 LinnaMis (and alunit the same time Lndwii:) used it in the sense of a proper calyx, yet \\iilisome vagueness. Mirliel and Brown established it in tlu1 sense of the collective floral eoverino;. DeCandolle revived Khrhart's FIG. 307. The complete ll.vwer .if a Crassula. 30S. Mia-rrain of its cross-section in the buil. showing the relative position of its parts The live pieces of the exterior circle a re sections of the sepals: the next, of the petals; the third, of the stamens through their :mlhcrs; the innermost, of the live pistils. KKi. 30! i A sep;il; 310, a petal; 311, a stamen; and 312, a pistil from the flower represented in Fig. 307. ous. ITS NATURE AND PARTS. 165 is as highly colored. A name being wanted for the individual leaves which make up the calyx, analogous to that for corolla- leaves, DeCandolle adopted Necker's coinage of the word sepal. Calyx-leaves are SEPALS. •JUS. The Corolla is the inner set of floral envelopes, usually (hut not always) of delicate texture and other than green color, form- ing therefore the most showy part of the blossom. Its several leaves are the PETALS.1 299. The floral envelopes are for the protection of the organs within, in the bud or sometimes afterward. Also, some of them, by their bright colors, their fragrance, and their saccharine or other secretions, serve for allurement of insects to the blossom, to mutual advantage. (504.) This furnishes a reason for neutral flowers, those devoid of essential organs, which sometimes occur along with less conspicuous perfect ones. "The leaves of the flower " are therefore indirectly subservient to reproduction. 300. The essential organs, being commonly plural in number, sometimes need a collective name. Where- fore, the aggregate stamens of a flower have been called the ANDRCECIUM ; the pistils, the GYNCECiUM.2 301. The Stamens3 are the male or fertilizing organs of a flower. A complete stamen (Fig. 311, 313) consists of FILAMENT (/), the stalk or support, and ANTHER (a), a double sac or body of two cells, side by side, filled with a powdery substance, POLLEN, which is at length discharged, usually through a slit or cleft of each cell. well-formed name of perifjonitim, and in the sense here given. But later (in the Organographie) he proposed to restrict it to cases in which the part is of ambiguous nature, as in Monocotyledons. The earlier definition is no doubt the proper one ; but the occasions for using the term in descriptive botany are mainly where the nature may seem to be ambiguous or con- fused, or where, from the union or close similarity of outer and innercircles, it is most convenient to treat the parts as forming one organ. 1 Fabius Columna, at the close of the sixteenth century, appears to have introduced this term, or, as Tournefort declares, " primus omnium quod sciam Petali vocem proprie usurpavit, ut folia florum a foliis proprie dictis distingueret." - The male household and the female household respectively, terms in- troduced by Keeper (Linnasa, i. 437), in the form of andrce.cev.rn a.ndgyna;ceum ; but the diphthong in the latter should also be , by Linn;ens named Germen), the hollow portion at the base which contains the a OVULES, or bodies destined to be- come seeds ; the STYLE (&), or colum- nar prolongation of the apex of the ovary ; and the STK.M \ (<•), a portion of the surface of the style denuded of epidermis, sometimes a mere point or a small knob at the apex of the style, but often forming a single or double line running down a part of its inner face, and assuming a great diversity of appearance in different plants. The ovary and the stigma are the essential parts. The style (as also the lilament of a stamen) may be altogether wanting. .31.5 apices. It came in time to be used as now for the whole organ ; but Lud- wig (List. Reg. Veg.), in 1741', apparently lirst so defined it, and introduced tlic term Anther for the A/n.r of Ray, or Th«-t pars interior et media floris, qua1 ex ovario et stylo com- ponitur. . . . Oriir/ni/i est pars pistilli inferior, qua- futuri fructus delinea- tioiiem sistit ^ti/litx est pars pistilli ex ovario centro produeta. . . • Summitas styli vel ejus partium S/i,/m,i dicitnr." Ludwiur. lust. Heg. Ycg. 41-43, 1742. Without mentioning the plural, the pistil is thus defined in a way which necessitates its use. Linna-us (in Phil. Bot.) first defines Stamen and 1'istillum in the singular number, enumerating the three parts of the latter, and afterwards (p. 57) declares that " Pistilla differimt quoad FIG. 314. Vertical section of a pistil, showing the interior of its ovary, a, to one side of which are attached numerous ovules, d: above is the style, b, tipped by the stigma, c. FIG. 315. A pistil of Crassula. like that of Fig. 312, but more magnified, and cut across through the ovary, to show its cell, and the ovules it contains; also pulled open below at the suture. At the summit of the style is seen a somewhat papillose portion, destitute of epidermis, extending a little way down the inner face: this is the stigma. ITS NATURE AND PARTS. 167 303. The Toms, or Receptacle of the flower, also named THALAMUs,1 is the axis which bears all the other parts, that upon which they are all (mediately or immediately) inserted. These are all ho- mologous with leaves. This is extremity of stem, or floral axis, out of which the organs described grow, in succession, like leaves on the stem ; the calyx from the very base, the petals .11 e next within or above the calyx, then the stamens, finally the pistils, which, whether several or only one, terminate or seem to terminate the axis. (Fig. 316.) 304. Metamorphosis. If flower-bnds are homologous with leaf-buds, and the parts of the flower therefore answer to leaves modified to special functions (293), then the kind of flower here employed in explaining and naming these parts is a proper pattern blossom. For the organs are all separate pieces, arranged on the receptacle as leaves are on the stem, the outer- most manifestly leaf-like, the next equally so in shape, though not in color, the stamens indeed have no such outward resem- blance, but the ripe pistils open down the inner angle and flatten out into a leaf-like form. The adopted theoiy supposes that stamens and pistils, as well as sepals and petals, arehomolo- numerum," etc., and so elsewhere, besides founding his orders on the num- ber of pistils. Among even French authors, Mirbel (1815) writes, "Le nombre des pistils n'est pas le meme dans toutes les especes," &c. Moquin- •Tandon freely refers to pistils in the plural, and Aug. St. Hilaire takes wholly the view here adopted, distinguishing the solitary pistil into simple and compound. DeCandolle, in Theorie Ele'mentaire, third edition, writes, " Chaque carpel est un petit tout, un pistil entier, compose d'un ovaire, d'un style, et d'un stigmate." Of English authors, no other need be cited than Robert Brown. The terms in question, then, are : — Gyncechtm, the female system of a flower, taken as a whole. Pistil, each separate member of the gyncecium ; this either simple or compound. Ovary, the ovuliferous portion of a pistil. Substituting a part for.the whole, this term is often used when the whole pistil is meant. Carpel, or Carpid, or Carpophytt, each pistil-leaf ; whether distinct as in simple or apocarpous pistils, or in combination of two or more to form a com- pound or syncarpous pistil. 1 By Tournefort, and adopted by Ludwig. Receptaculum floris, Linnaeus. Thorns, Salisbury. Torus (the proper form), DeCandolle. FIG. 316. Parts of the flower of a Stonecrop, Sedum ternatum, two of each sort, and the receptacle, displayed-, a, sepal; b, petal; c, stamen; rf, pistil. 168 THE FLOWER. gous with leaves ; that the sepals are comparatively little, the petals more, and the reproductive organs much modified i'roin the type, that is from the leaf of vegetation. This is simply what is meant hy the proposition that all these organs are transformed or metamorphosed leaves. What would have been leaves, if the development had -one on as a vegetative branch, have in the blossom developed iii other forms, adapted to other func- tions. Limueus expressed this idea, along with other more speculative conceptions, dimly apprehended, by the phrase Vege- talile .Metamorphosis. Not long afterwards, this fecund idea of a common type, the leaf, of which the parts of the flower, &c., were regarded as modifications, was more clearly and differ- ently developed by a philosophical physiologist, Caspar Frederic Wolff. Thirty years later, it was again and wholly independ- ently developed by Grcthe, in a long-neglected but now well- known essay, on the Metamorphosis of Plants. Twenty-three years afterwards, similar ideas were again independently pro- pounded by DeCandolle, from a different theoretical point of view; and finally the investigation of phyllotaxy has completed the evidence of the morphological unity of foliaceons and floral organs.1 1 The contribution of Linnaeus is on p. 301 of the Philosophia Botanica, 1751 ; and all that is pertinent is in the following propositions : — I'lumulam seminis saepius terminal aut flos aut gemma. Principium florum et foliorum idem est. Prindpiuni gemmarum et foliorum idem est. Gemma constat foliorum rudiment is. Periantliium sit ex connatis foliorum rudimcntis. His dissertation, Prolepsis I'lantarum, in Anwn. Acad. vi. (1700), added nothing hut obscure speculations to the former comparatively clear statements. Kaspar Fricdrich Wolff's contribution is in his Theoria Generationis, mainly concerning animals, published in 1750, and an enlarged and amended edition in 1774. He first clearly conceives the plant as formed of two ele- ments, stem and leaf, but develops only the morphology of the latter, and under the hypothesis that leaves of vegetation become bud-M-iles or tloral organs, as the ease may he, through degciiercscence or diminution of vege- tative force, which is renewed in the hud or in the seed. Johann Wolfgang (Jothe's Yersneh die Metamorphose der Prlan/cn zu erkliiren was published in 17'.M). in 8(5 pages. For the translations and reproduction.-, see Prit/.el, Thesaurus. To tin- Kreiich translation by Soret. with (Jerman text accompanying (Stuttgart, ls:!l), and a No to that of Ch. Martens |(Kuv. I list. Na t. de ( iirthe, Paris, 1837), are joined the author's inter- esting notes and anecdote- of later periods, down to 1 S-">1. The degcncrescence by diminution of vegetative force with renewals by generation, propounded by Wolff, in (in tlie's essay takes the form of successive expansion and con- traction of organs. A. P. DeCandolle's Theorie El&nentaire de Botanique appeared in 1813, ITS METAMORPHY. 169 305. It will be understood that metamorphosis, as applied to leaves and the like, is a figurative expression, adding nothing to our knowledge nor to clearness of expression, but rather liable to mislead. The substance of the doctrine is unity of type. Its proof and its value lie in the satisfactory explanation of the facts, all of which it co-ordinates readily into a consistent and simple system. As applied to the flower, two kinds of evidence may be adduced, one from the normal, the other from teratological conditions of blossoms. The principal evidence of the first class is that supplied by 306. Position and Transitions. As illustrated in the preced- ing chapter, the flower occupies the place of an ordinary bud or leaf-bud. Also the parts of the flower are arranged on the receptacle as leaves are arranged on the stem, i. e. they conform to phyllotax}7, as well in passing from leaves and bracts to the perianth, as in the position of the floral organs in respect to each other. This is partly shown in the preceding chapters, and is to be further illustrated. Sepals, petals, stamens, and pistils are either in whorls or in spirals, and have nothing in their arrangements as to position which is not paralleled in the foliage. 307. The evidence from transitions has to be gathered from a great variety of plants. Very commonly the change is abrupt from foliage to bracts, from bracts to calyx -leaves, from these to corolla-leaves, and from these to stamens. But instances abound in which every one of the intervals is bridged b}' transitions or a second edition 1819 ; a third (revised by Alphonse DeCandolle), in 1844, is posthumous. The Organographie Vegetale, in which the morphology of the earlier work is developed, appeared in 1827. The leading idea is that of symmetry, of organs symmetrically disposed around an axis (the homology of foliar and floral organs not at first apprehended), but this symmetry disguised or deranged more or less by unions (solderings) of homogeneous or heterogeneous parts, by irregularities or inequalities of growth, by abortions, &c. The reason why the organs in question have a normal symmetrical dis- position on the vegetative and floral axes was not reached by DeCandolle, nor was it perceived that the arrangement of leaves and of floral organs was identical. All this was the contribution of phyllotaxy, — a subject which was approached by Bonnet (an associate of DeCandolle's father), and first investigated by the late Karl Schimper and Alexander Braun, beginning about the year 1829. It is interesting to know that Wolff's work was wholly unknown to Gcethe in 1700, and that both Wolff's and Goethe's were unknown to DeCan- dolle until after the publication of the second edition of the latter's The'orie Elementaire, in 1819. When the Organographie appeared, the essay of Goethe had come to light ; and contemporary contributions to floral morphology by Petit-Thouars, R. Brown, Dunal, and Rceper, were adding their influence. 170 THE FLOWER. intermediate forms. The gradual transition from ordinary foli- age to bracts and bractlets is exceedingly common. In color and texture it is not rare to meet with bracts which vie , with, or indeed surpass, pet- als themselves in delicacy and brightness; and in such cases they assume a principal oilire of flower-leaves, that of con- spicuous show for attraction. Scarlet Sage, Painted-Cup (Castilleia), and the I'oin- settia. with other Euphorbias of the conservatories, are ex- amples of this. In the (lowers of Barberry, it is by a nearly arbitrary selection that bractlets are distinguished from sepals : in Calycanthus, in many kinds of Cactus, and in Nelumbium, the same is true as to bractlets, se- pals, and petals ; in Water-Lily (Nym- plitva, Fig. :'>!*). there is a gradual transition from the sepals through the IIJI /fll petals to stamens : rfc ' //// // //a in Lilies and most // / // 1 /f*/ lily-like flowers, se- \ If pals are as brightly (I colored as petals, and commonly more or less combined with them. When the perianth-leaves are of only one set, it is not at all by color or texture that this perianth can be assigned to calyx or to corolla. Normal transitions from a stamen to a pistil could not, in the nature of the case, he expected. .".os. Tcratolofrical Transitions aiid Changes. Teratology is the study of monstrosities. These in the vegetable kingdom 818 FIG. 317. Cactus-flower (Mamillaria caespitosa), with bractlets, sepals, and petals passing into o.-icli other. FK J. 318. Series exhibiting transition from sepals to stamens in Nymphsea odorata. ITS METAMORPHY. 171 often elucidate the nature of organs.1 The commonest of these changes belong to what was termed by Grethe retrograde meta- morphosis ; that is, to reversion from a higher to a lower form, as of an organ proper to the summit or centre of the floral axis into one which belongs lower down.2 The most familiar of all such cases is that of the so-called double flower •, better named in Latin flos plenus. In this, the essential organs, or a part of them, are changed into colored flower-leaves or petals. Most flowers are subject to this change under long cultivation (witness " double" roses, camellias, and buttercups), at least those with numerous stamens. It occasionally occurs in a state of nature. The stamens diminish as the supernumerary petals increase in number ; and the various bodies that may be often observed, inter- mediate between perfect £ stamens (if any remain) and the outer row of petals, — from imperfect petals, with a small lamina tapering into a slender stalk, to those which bear a small distorted lamina on one side and a half-formed anther on the other, — plainly reveal the nature of the transformation 319 that has taken place. Carried a step farther, the pistils likewise disappear, to be replaced by a rosette of petals, as in fully double 1 The leading treatises are Moquin-Tandon's Teratologie Ve'ge'tale, Paris, 1841, and Masters, Vegetable Teratology, London, published for the Ray Society, 1869. An earlier publication deserves particular mention, viz. the thesis De Antholysi Prodromus, by Dr. George Engelniann, Frankfort on the Main, 1832. 2 To these abnormal changes, the term metamorphosis is obviously more applicable ; for here what evidently should be stamens, pistils, &c., on the testimony of position and the whole economy of the blossom, actually ap- pear in the form of some other organ : yet even here the change is only in the nisus format ivus ; the organ was not first formed as a stamen, and then transformed into a petal or leaf. FIG. 319. A flower of the common "White Clover reverting to a leafy branch ; after Turpin. Calyx with tube little changed, but lobes bearing leaflets. Pistil stalked ; the ovary open rlown the inner edge, and the margins of the pistil-leaf bearing leaves instead of ovules. 172 THE FLOWER. buttercups.1 In these the green hue of the centre of the rosette indicates a tendency to retrograde a step farther into sepals, or into :i duster of green leaves. This takes place in certain blos- soms of the Strawberry, the Rose, &c. Such production of ••"Teen roses," and the like, has been appropriately called chlorosis, or by Masters r/,A,/v,/,%. from the change to irreen. ;;ui>. A monstrosity of the blossom of White Clover, long ago figured by Turpin (Fig. 319), is such a case of foliaceous rever- sion, in which even the ovules are implicated. The imperfect leaves which take the place of the latter may be compared with the leaty tufts which form along the margins of a leaf of Bryophyllum, by which the plant is often propagated. (Fig. 322.) 310. The reversion of a simple pistil di- rectly to a leaf is seen in the Double-flowering Cherry of cultivation (Fig. 320, 321), usually passing moreover, by prolificatiou of the re- ceptacle, into a leafy branch. 311. The reversion of pistils to stamens is rarer, but has been observed in a good number of instances, in Chives, in the Horseradish, in Gentians and Hyacinths, and in some Willows. In the latter, the opposite transformation, of stamens to carpels, is vei-v common, and curious grades between the two are met with almost every spring. So also in the common llouseleek, and in perennial Larkspurs. Certain apple-trees are known, both in the United States and Europe, in which, while the petals are changed into the appearance of minute green sepals, the outer stamens are converted into carpels, these supernumerary and in the fruit superposed to the five normal carpels.- In Poppies, many of the innermost stamens are occasionally transformed into as many small and stalked simple pistils, surrounding the base of the large compound one. 1 It must not be conclude.! that the supernumerary petals in all such cases are reverted stamens, or stamens and pistils. Sonic are instances of abnormal plciutaxy, /. .•. of the production of our or more additional ranks of petals (better deserving the name of ilonlilc jion-i r), with or without reversion of essential organs to flower leaves. 2 These trees are popularly supposed to hear fruit without blossoming; the reverted green petals being so inconspicuous that the tlower is un- noticed. FIG. 320. 321. Green loaves from the centre of a blossom of Double-flowering Cherry, one still showing. l>y its partial involution anil its style-like apex, that it is a reverted carpel, the other a small but woll-fnnneil leaf. FIG. 322. Leaf ur leallet of Hryophyllum, developing plantlets along the margins. ITS METAMORPHY. 173 312. Another line of teratological evidence is furnished by prolification. The parts of the flower are, by the doctrine, homologous with leaves, and no leaf ever terminates an axis. Normally, in fact, the axis is never prolonged beyond the flower, but abnormally it ma}' be1. It may resume vegeta- tive growth as a termi- nal growing bud, either from between the pistils after the whole flower is formed, or at an earlier period, usurping the central part of the flower. Thus, when a rose is borne on a pe- duncle rising from the centre of a rose, which is not very unusual, or a leafy stem from the top of a pear (Fig. 323), the flower was probably complete before the monstrous growth set in. In Fig. 324, the reversion to foliaceous growth took effect after the stamens but before the pistils were formed. In rose-buds out of roses, the terminal proliferous shoot takes at once the form of a peduncle ; in the shoot from the pear, that of a leaf}7 stem. 313. Again, axillary buds are normally formed in the axil of leaves. No such branching is known in a normal flower. But in rare monstrosities a bud (mostly a flower-bud) makes its appearance in the axil of a petal or of a stamen ; and it may be clearly inferred that the organ (not itself axillary) from the axil of / which a bud develops is a leaf or its N homologue. Fig. 325 exhibits a clear case of the kind, a flower in the axil of each petal of Celastrus scandens. Flowers, or pedunculate clusters of flowers, from the axil of petals of garden Pinks are sometimes seen. A long-pedunculate flower from the axil of a FIG. 323. A monstrous pear, prolonged into a leafy branch ; from Bonnet. FIG. 324. Retrograde metamorphosis of a flower of the Fraxinella of the gardens, from Lindley's Theory of Horticulture ; an internode elongated just above the stamens, and hearing a whorl of green leaves. FIG. 325. A flower of False Bittersweet (Celastrus scandens) producing other flowers in the axils of the petals ; from Turpin. 174 THE FLOWER. stamen of a species of "Water-Lily (Nymphcea Lotus) is figured and described l>y Dr. Masters.1 314. In the application of morphological ideas to the elucida- tion of the flower, nothing should be assumed in regard t<> it which has not its proper counterpart and exemplar in the leaves and axis of vegetation. SECTION II. FLORAL SYMMETRY. 315. The parts of a flower are symmetrically arranged around its axis.2 Even when this symmetry is incomplete or imperfect, it is still almost always discernible ; and the particular numerical plan of the blossom may be observed or ascertained in some of the organs. 31G. Adopting the doctrine that the parts of the flower are homologous with leaves, the symmetry is a consequence of the phyllotaxy. It is symmetry around an axis, not the bilateral symmetry which prevails in the animal kingdom. For parts of a flower disposed in a continuous spiral (which mostly occurs when they are numerous), the arrangement is that of some order of this kind of phyllotaxy, which distributes the parts equably into superposed ranks. (237.) The much commoner case of 1 The fullest enumeration and discussion of the very various kinds of abnormal structures and deviations in plants is to be found in the Teratology of Dr. Masters, above referred to. Many technical terms are here brought into use, which need not be here mentioned, except the following, which relate directly to floral metamorphosis. /'//////-/(/// (called Phyllomorphy by Morren. /'/••<» affected. J'ixtil/ix/i/. where other organs develop into pistils, which most rarely happens except with the .stamens, as abo\e mentioned. : It is stated that ( 'orrea de Serra (who published botanical and other papers in London. Paris, and Philadelphia during the first twenty years of the century, but who knew far more than he published) was the first botanist to insist on the symmetry of the flower. It was first made prominent by De Candolle, in the Theoric Elementaire, and elaborated in detail by A. St. llilaire in his Morphologic Ve'ge'tale. FLORAL SYMMETRY. 175 equal number of parts in a cycle, and the cycles alternating with each other, is simply that of verticillate phyllotaxy. (234.) In either case, the members of the successive circles (or of closed spirals as the case may be) will be equal in number ; that is, the flower will be isomerous. 317. A Symmetrical Flower is one in which the members of all the cycles (whorls or seeming whorls) are of the same number.1 In nature, the symmetry is of all degrees : it is most commonly complete and perfect as to the floral envelopes when it is not so as respects the essential organs. The general rule is that the successive cycles alternate, as is the nature of true whorls. But the superposition of successive parts is not incompatible with symmetry of the blossom, although it is a departure from the ordinary condition, assumed by botanists as the type. An isomerous flower (meaning one with an equal number of mem- bers of all organs) is the same as sjonmetrical, if the reference be to the number in the circles, rather than to the total number of organs of each kind. 318. A Regular Flower is one which is symmetrical in respect to the form of the members of each circle, whatever be their number ; ?'. e., with the members of each circle all alike in shape. 319. These two kinds of symmetry or regularity, with their opposites or departures from symmetry, need to be practically distinguished in succinct language. For the terminology, it is best to retain the earlier use, generally well established in phyto- graphy, as above denned. 320. A Complete Flower is one which comprises all four or- gans, viz. calyx, corolla, stamens, pistil. 1 This is not only the definition " generally applied in English text-books," but that introduced by DeCandolle, adopted by St. Hilaire, and followed at least by the French botanists generally. The innovating German definition, of a recent date, is that a symmetrical flower is one " that can be vertically divided into two halves each of which is an exact reflex image of the other." But such have immediately to be distinguished into " flowers which can be divided in this manner by only one plane," which Sachs terms " simj>/i/ symmetrical or monosymmetrical," and those which can be symmetrically divided by two or more planes, " doubly symmetrical or poll/symmetrical," as the case may be. Now both these forms have a more expressive and older terminology, adopted by Eichler, viz. : — Zygomorphous, for flowers, or other structures, which can be bisected in one plane, and only one, into similar halves (median zygomorphous, when this is a median or antero-posterior plane, as it most commonly is ; trans- verse zygomorphous, when the plane of section is transverse or at right angles to the median, as in Dicentra) ; Actinomorphous for flowers, &c., which can be bisected in two or more planes into similar halves. 176 THE FLOWER. 321. Numerical ground-plan. Many flowers are numerically indefinite in some or most of their kinds of members, as Ranun- culus, Magnolia, and the Rose for stamens and pistils, Xym- phsea for all but perhaps the sepals, man}- Caetacca' for all but the pistil, and Calycanthus for all four components. But more commonly each flower is constructed upon a definite numerical ground-plan ; and the number is usually low. Seldom, if ever, is it reduced to unity in a hermaphrodite blossom (even Ilip- puris, with a single stamen and a single pistil, is not an un- equivocal case), and probably never in a complete one. But there are such extremely simplified flowers among those of a single sex. In Monocotyledons, the almost universal number is three, some- times two ; in ordinary Dicotyledons, five prevails ; four and two are not uncommon ; three is occasional ; and higher numbers are not wanting, as twelve or more in Houseleeks. 322. To designate the particular plan, such familiar terms of Latin derivation as binary, ternary, i/nati'nian/. quinary, senary. &c., are sometimes employed, denoting that the parts of the flower are in twos, threes, fours, fives, or sixes. More technical and precise terms, equivalent to these, are composed of the Greek numerals prefixed to the word meaning parts or members, as Monomer ous, for the case of a flower of one member of each ; IHmerous, of two, or on the plan of two members of each ; Trimerous, of three, or on the plan of three members ; Tetramerous, of four, or on the quaternary plan ; Pentamerous, of five, or on the quinary plan ; Hexamerous.1 of six, or on the plan of six members to each circle. But, in Monocotyledons, so-called hexamerous blossoms are really trimeroiis, the sixes being double sets of three. Pattern Flowers. These should be >•////////• tri<-H. may serve as a pattern pentamerous or quinary flower: and Fig. 3i>r>. with its diagram. Fig. 327, These may be shortly written 1-inerous, 2-iiien>u>. .'1-iniToiis, anil so on up in 10-merouB (decamerous), 1'J-merous (dodecamerous), &c. Fit;. :;L'I;. Parts nf a symmetrical trimcnms ilowrr iTilla.-a muscosa): a. calyx; 6 corolla; r. stamens; d. pistils 327. Diagram of tbe same. FLORAL SYMMETRY. 177 as a pattern trimerous or ternary flower; these being simply isomerous, and of one circle of each kind. And the whole relation of the parts, viewed as modified leaves on the common axis, may be exhibited in such a diagram of a pattern isostemonous 5-merous flower as that displayed in Fig. 328. 324. Diplostemonons Type. The foregoing patterns are selected upon the idea of the greatest simplicity consistent with com- pleteness. But extended observation leads to the conclusion that the typical flower in nature has two series of stamens, as it has two series in the perianth ; that is, as many stamens as petals and sepals taken together.1 As the petals alternate with the sepals, so the first series of stamens al- ternates with the pet- als, the second series of stamens alternates with the first, and the pistils or carpels when of the same number alternate with these. Thus the outer series of stamens and the carpels normally stand before (are 1 This view of the symmetry of the flower was first taken by Brown (Obs. PI. Oudney, in Denham and Clapperton Trav. 1826, reprinted in Ray Soc. ed. of Collected Works, i. 293). It is true that Brown declares the same of the pistils ; but that is not made out. The evidence of this doctrine is to be gathered from a large and varied induction ; from the general pres- ence of the two sets of stamens, and no more, in petaloideous Monocotyle- dons ; the unaltered position of the carpels (before the sepals) when the inner set of stamens is wanting, as in the Iris Family ; the very common appearance in haplostemonous flowers among the Dicotyledons of vestiges of a second series, or of bodies which may be so interpreted. The andrcecium or the blossom is said to be Isostemonous or Haplostemonous when the stamens are of one series, equal in number to that of the ground-plan of the blossom ; Diplostemonous, when there are two series, or double this number. FIG. 328. Ideal plan of a plant, with the simple stem terminated by a symmetrical pentamerons flower; the different sets of organs separated to some distance from each other, to show the relative situation of the parts. One of each, namely, a, a sepal, b. a petal, c, a stamen, and d, a pistil, also shown, enlarged. FIG. 329. A pentamerous diplostemonous flower of Sedum. ITS THE FLOWER. superposed to) the sepals, and the stamens of the inner series stand before the petals ; as in the diagram. Fiji. 331.1 :'>'2~>. Flowers which completely exemplify their type or symmetry are rare, but mo>t exhibit it more or less. Kadi natural order or group exhibits its own particular floral type, or modification of the common type.- Some of these modifications do not at all affect the sy linnet ry or obscure the plan of the flower, except by combina- tions which render the phylline character of the floral envelopes and carpels less apparent, such combinations being of rare occurrence in foliage. Others gravely interfere with floral symmetry, sometimes to such degree that the true plan of the blossom is to be ascertained only through extended comparisons with the flowers of other plant* of the same order or tribe, or of related orders. The symmetiy of the blossom finds its explanation in the laws which govern the arrangement of leaves on the axis; that is, in phyllotaxy. The deviations from symmetry and from typical simplicity have to be explained, and in the first instance 1 For convenient reference and the avoidance of circumlocution, some \\ritiTs term the stamens which are before tin- petals epipefo/ous, those before the sepals , /lisr/mlnu* ; but, as this prefix means upou, it is better to restrict these terms to rases of adnation of stamens to these respective parts of the perianth, and to distinguish as Anii/K /n/inis, those stamens which stand before petals, whether adnate or free, and Antisepalous, those which stand liet'oiv sepals. — These terms we find have already been employed in this way by Dr. A. Dickson (in Seemann, Jour. Bot. iv. 275), with the addition of a third, viz. /'iini/Kfii/tiiia, for stamens which stand at each side of a petal, yet not necessarily before a .sepal, as in many Rosaceae. 2 These particular types, with their modifications, are set forth in the i-fiiirin-i, rs or distinguishing marks of the orders, tribes, genera, Xc. The best generally available illustrations of ordinal types are in Le Maoiit and Decaisne's Traite (ie'ne'ral de Botaniqiie, and in Hooker's Knglisli edition and revision, entitled A (ieneral System of Botany, Descriptive and Analyti- cal, London, 187'J. The best morphological presentation is in Eichler's Bluthendiagramme, £c. (Flower Diagrams, Constructed and Illustrated), Leipzig, 1875. FIG. 330. Opened flower of Trillium erectum. 331. Diagram of the same. ITS GENERAL MODIFICATIONS. 179 to be classified. To have morphological value, such explanation should be based upon just analogies in the foliage and other organs of vegetation. Whatever is true of leaves and of the vegetating axis as to position of parts, mode of origin and growth, division, connection, and the like, may well be true of homologous organs in the flower. SECTION III. VARIOUS MODIFICATIONS OF THE FLOWER. § 1. ENUMERATION OF THE KINDS. 326. In the morphological study of flowers, these modifica- tions are viewed as deviations from type. Their interpretation forms no small part of the botanist's work. They may be classed under the following heads : - 1. Union of members of the same circle : COALESCENCE. 2. Union of contiguous parts of different circles : ADNATION. 3. Inequality in size, shape, or union of members of the same circle : IRREGULARITY. 4. Non-appearance of some parts which are supposed in the type : ABORTION or SUPPRESSION. 5. Non-alternation of the members of contiguous circles : ANTEPOSITION or SUPERPOSITION. 6. Increased number of organs, either of whole circles or parts of circles : AUGMENTATION or MULTIPLICATION. 7. Outgrowths, mostly from the anterior or sometimes pos- terior face of organs : ENATION. 8. Unusual development of the torus or flower-axis. 9. To which may be appended morphological modifications, some referable to these heads and some not so, which are in special relation to the act of fertilization. These are specially considered in Section IV. 327. These deviations from assumed pattern are seldom single ; possibly all may coexist in the same blossom. Several of them occur even in that one of the orders, the Crassulacese, which most obviously exhibits the normal type throughout. 328. Thus, Sedum (Fig. 329), with two circles of stamens, being taken as the true type (324), Crassula (Fig. 307) wants the circle of stamens before the petals ; Tillsea (Fig. 326) is the same, but with the members S3Tnrnetrically reduced from five to three ; Rhodiola loses all the stamens by abortion in one half the individuals and the pistils in the other, sterile rudiments testifying to the abortion ; Triactina has lost two of its five carpels, and t^e three remaining coalesce into one body up to the 180 THE FLOWER. middle ; Penthorum (Fig. 335, 336) has its five carpels coales- v Kimia-us to an occasional monostrosity of these flowers (iniitatcil in sundry others), in which tin- base of every petal, or answering part of tin- corolla, is prolonged downward into a sac or spur. The sac is, morphologically considered, a depart ure from normal regularity : in the monster, symmetrical regularity is restored by the development of four more sacs. FIG :M!>. Flower cif Viola sagittata. 350. Its sepals anil petals displayed. 351. Diatrr.-im nf :i Yiol.-t-Mossoin. from KMiler, with bract orsulitrn 'lint; leaf (below), a pair of bractlets (lateral), ami axis to which the subtending leaf belongs (above or posterior). DISAPPEARANCE OF PARTS. 187 § 5. DISAPPEARANCE OR OBLITERATION OF PARTS. 342. Abortion or Suppression are somewhat synonymous terms to denote the obliteration or rather non-appearance of organs which belong to the plan of the blossom. Abortion is applied particularly and more properly to partial obliteration, as where a stamen is reduced to a naked filament, or to a mere rudiment or vestige, answering to a stamen and occupying the place of one, but incapable of performing its office ; suppression, to abso- lute non-appearance. Such vestiges or abortive organs justify the use of these terms, the more so as all gradations are some- times met with between the perfect organ and the functionless rudiment which occupies its place. Such obliterations, whether partial or complete, may affect either a whole circle of organs or merely some of its members. The former interferes with the completeness of a flower, and may obscure the normal order of its parts. The latter directly interferes with the symmetry of the blossom, and is commonly associated with irregularity. 343. Of parts of a Circle. Among papilionaceous flowers (338) , different species of Erythrina have all the petals but one (the vexillum, Fig. 344, a) much reduced in size, in some concealed in the calyx, and in every way to be ranked as abortive oi'gans. In Amorpha, of the same family, these four petals are gone, leaving no trace, reducing the corolla to a single petal. (Fig. 352, 353.) This one is evidently the vexillum, both by position and shape ; and the 5-merous type, also the particular type of the family, are still discernible in the five notches of the calyx, the ten stamens, &c. In a related genus, Parryella, even this last petal is wanting, and the androecium is straight, all irregularity thus disappearing through suppression. 344. Delphinium or Larkspur and Aconite or Monkshood furnish good examples of flowers in which irregularity is accom- panied by more or less abortion. The calyx of the Larkspur (Fig. 354-356) is irregular b}' reason of the dissimilarity of the five sepals, one of which, the uppermost and largest, is pro- longed posteriorly into a long and hollow spur. Within these, and alternate with them as far as they go, are the petals, only FIG. 352. Stamens and pistil of Amorpha fruticosa. 353. An entire flower of the same. 18S THE FLOWER. lour in nuinbor, and those of two shapes, the two upper ones having long spurs which are received into the spur of the upper sepal ; the two lateral ones having a small but broad blade raised on a stalk-like claw ; and the place which the fifth and lower petal should occupy (marked in the ground-plan. Fig. 356, FIG. 354. Flower of a larkspur. :ir.r>. The five sepals (outer circle) and the four petals (inner circle) displayed. 350. Ground-plan of tin- calyx and corolla. FIG. :t.r>7. Flower of an Aconite or Monkshood. 35S. The fm- sepals and the two small and curiously shaped petals displayed; also the stamens and pistils in the centre. 359. Ground-plan of the calyx and corolla; the dotted lines, as in Fig. 356, representing the suppressed parts. DISAPPEARANCE OF PARTS. 189 by a short clotted line) is vacant, this petal being suppressed, thereby rendering the blossom unsymmetrical. In Aconite (Fig. 357-359), the plan of the blossom is the same, but the uppermost and largest of the five dissimilar sepals forms a helmet-shaped or hood-like body ; three of the petals are wanting altogether (their places are shown by the dotted lines in the ground- plan, Fig. 359) ; and the two upper ones, which ex- tend under the hood, are so reduced in size and so anomalous in shape that they would not be recog- nized as petals. One of these, enlarged, is exhibited in Fig. 360. Petals and other parts of this and of va- rious extraordinary forms were termed by Linnaeus NECTARIES, a somewhat misleading name, as they are no more devoted to the secretion of nectar than ordinary petals or other parts are. In these flowers, moreover, the stamens are much increased in number. 345. Analogous abortion of some of the stamens,' along with a particular irregularit}" of the perianth, especially of the corolla, characterizes a series of natural orders with coalescent petals.1 These flowers are all on the 5-merous plan (except that the gynoecium is 2-merous), but with corolla, and not rarely the calyx, irregular through unequal union in what is called the bila- biate or two-lipped manner. The greater union is always median, or anterior and posterior, and two of the coalescent members form one lip, three the other. The two posterior petals form the upper lip, the anterior and two lateral form the lower lip of the corolla ; in the calyx, when that is bilabiate, this is of course reversed. In some, as in Sage and Snapdragon, the bilabiation of the corolla is striking (Fig. 479-481), and readily comparable to the two jaws of an animal ; in others, the parts are almost regu- lar. The suppression referred to is, in most of these cases, that of the posterior of the five stamens, as in Fig. 361, where it is complete. In Pentstemon (Fig. 362), a sterile filament regu- larly occupies the place of the missing stamen. The position sufficiently indicates its nature. This is also revealed by the rare occurrence of an imperfect or of a perfect anther on this 1 These natural orders in which this occurs, or tends to occur, are the Scroplmlariaceae (Snapdragon, Pentstemon, Mimulus, £c.), Orobanchacese (Beech-drops), Lentibulaceae (Bladderwort), Gesneraceae (Gloxinia), Big- noniacese (Trumpet Creeper, Catalpa), Pedaliaceae (Martynia), Acanthacea?, Labiatae (Salvia, Stachys), &c. FIG. 360. A petal (nectary) of an Aconite, much enlarged. 190 THE FLOWER. filament, — a monstrosity, indeed, but the monstrosity- is here a return to normal symmetry. The two stamens nearest the suppressed or abortive one generally share in the tendency to abortion, as is shown by their lesser length or smaller anthers : in the flower of Catalpa, these two also are either im- perfect or reduced to mere vestiges (as in Fig. 363) : in very many other plants of these families, even these vestiges are not seen, and so the five stamens are by abortion or complete suppression reduced to two. 346. Suppression in the gynoecium to a number less than the numerical plan of the flower (as shown in the perianth) is of more common occur- rence than the typical number, and the reduction is comparatively con- stant throughout the genus or order. A papilionaceous or other leguminous flower with more than one or with all five pistils is exceedingly rare, and except in one pentacarpellaiy genus is a monstrosit}'. Suppression of the interior is more common than of exterior organs. Want of room in the bud may partially explain this. 347. Suppression of whole Circles. Such suppression or rather non-production in the actual blossom of whole series of organs which belong to tin- type, and indeed are sometimes present in that blo-^i MII'S nearest relatives, is very common. It gives occasion to several descriptive terms, which may be here defined together. First, and in general, flowers are liK-niii/ilefe, in which any one or more of the four kinds of organs is wanting, whatever these may be : .\/n't(i/<>iis. when the corolla or inner perianth is wanting; Monochliinn/ilcons, where the perianth is simple instead of FKl. :.«;i. Corolla of i ierardia purpurea laid open, with the four stamens, the place which the fifth should occupy indicated hy a cross. FI(!. .'ii'.'j. ( 'orolia of IVnistrinoii craiiditlorus laid open, with its four stamens, and a sterile lilanient in the place of the lifih stamen. Kit;. 363. Corolla of Catalpa laid open, with two perfect stamens and the vestiges of throe abortive ones. DISAPPEARANCE OF PAKTS. 191 double, in which case the wanting set is generally (but not quite alwa^'s) the inner, or the corolla ; Dichlamydeous, when both circles of the perianth (calyx and corolla) are present ; Achlamydeous, when both are wanting, as in Fig. 365. (These three terms are seldom employed.) Unisexual (also Diclinous or Separated) , when the suppression is either of the stamens or the pistils. In contradistinction, a flower which possesses both is Bisexual or Hermaphrodite. Staminate, or Male, when the stamens are present and the pistils absent ; Pistillate, or Female, when the pistils are .present and the stamens absent ; Monoecious (of one household), when stamens and pistils oc- cupy different flowers on the same plant ; Dioecious (of two households) , when they occupy different flowers on different plants ; Polygamous, when the same species bears both unisexual and bisexual or hermaphrodite flowers. This majr occur in various wa}'S, from the greater or less abortion of either sex, either on the same or on separate individual plants ; — as Monceciously or Diceciously Polygamous, according to the tendency to become either monoscious or dioecious. Recently Darwin has well distinguished the case of Gyno-dicecious, where the flowers on separate individuals are some hermaphrodite and some female, but none male only ; and Andro-dicecious, of hermaphrodite flowers and male, but no separate female. The latter is a less common case. Neutral, as applied to a flower, denotes that both stamens and pistils are wanting, — a case neither rare nor inexplicable on grounds of utility. (356, 504.) Sterile and Fertile are more loosely used terms. A sterile flower may mean one which fails to produce seed, as a sterile stamen denotes one which produces no good pollen, and a sterile pistil one which is incapable of seeding. But commonly a sterile flower denotes a staminate one ; a fertile flower, one which is pistillate, if not also hermaphrodite. 348. Suppressed Perianth. Almost universally, when the peri- anth is reduced to a single circle, it is the inner, or corolla, which is not produced. Or, rather, when there is only one circle or sort of perianth-leaves, it is called calyx, whatever be the appearance, texture, or color, unless it can somehow be shown that an outer circle is suppressed. For since the calyx is frequently delicate and petal-like (in botanical language, petaloid or colored, as in 192 THE FLOWER. Clematis and Anemone, Fig. 364), and the corolla is sometimes greenish or leaf-like, the only real dilierenee between the two is that the calyx represents the outer and the corolla the inner series. Even this distinction becomes arbi- trary when the perianth consists of three or four circles, or of a Less definite number of spirally arranged members. :>4!». Vet the only perianth obvi- ously present may be corolla, as when the calyx has its tube wholly adnate to the ovary and its border or lobes obsolete or wanting.'1 Aralia nudie.-mlis ( Fig. 341) is an instance, likewise many Umhc-llifcrjB, some species of Feclin or Valerianella. the fertile flowers of Nyssa, and those Composite which have no pappus. For l'.\n>t;.s, the name originally given to thistle-down and the like, answers to the border or lobes of a calyx attenuated and depauperated down to mere fibres, bristles, or hairs. The name is ex- tended to other and less obliterated forms. (C.44, Fig. 631-633.) When the obliteration is complete, as in Mayweed (Fig. 630), in some species of Coreopsis. &<-.. the corolla seems to be simply continuous with the apex of the ovary. A comparison with related forms reveals the real state of things.2 O 350. So also in llippuris. in which (along with extreme numerical reduction of the other floral circles) the calyx as well as corolla seems to be wanting: but the insertion of the stamen on the ovary (epigynous) suggests an adnate ealyx, and near inspection detects its border. 351. Both calyx and corolla an- really want- ing in the otherwise complete and perfect (symmetrical and 1 In tin- flowers of the two common species of Prickly Ash (Zanthoxy- luni) of the Atlantic; United States. OIK- has a double, the oilier a single perianth (as shown in Cray. (ien. lllustr. ii. 148, t. lAli) : the position of the stamens gives a presumption that the missing circle of the latter is the calyx ; yet it may lie otherwise explained. In Santalaeea- there are some grounds for suspecting that the simple perianth, although opposite the stamens, is corolla : and the foliaceous sepal-lohes of the female (lowers of Buckleya would confirm this, if these are true sepals rather than adnate braets. J In the pappus of Composite, every gradation is seen between undoubted calyx, recognizable as such by structure as well as position, and diaphanous scales, bristles, and mere hairs, wholly " trichomes " as to structure, although in the place of "phyllomes " and representing them. Flower of Anemone Penns\ Ivanira; apetalous. tlie calyx petalni I. Hi ; :;uri. Aclilainyilenus flower of Lizard's-tail (Saururus cernuus), magnlried. DISAPPEARANCE OF PARTS. 193 trimerous) flowers of Saururus, Fig. 365. But achlamydeous blossoms are usually still further reduced to a single sex. 352. Suppression of one circle of stamens is of very common occurrence. It is seen in different species of Flax ; which have mostly 5-merous perfectly symmetrical and complete flowers with one set of stamens abortive. In some species (as in Fig. 3G7), 367 vestiges of the missing circle of stamens are conspicuous in the form of abortive filaments, interposed between the perfect stamens ; in others, these rudiments are inconspicuous or even altogether wanting. 353. Suppressed Androecium or Gynoecinm. This occurs with- out or along with suppression in the perianth. In cases of the former, vestiges of s;o sn the aborted organs often remain to sig- nify the exact nature of the loss. Sepa- ration of the sexes (monoecious, dioeci- ous, &c.) is the re- sult of sueh suppres- sion. In Menisper- mum (Fig. 368, 369), this is accompanied by an actual doub- ling of both calyx and corolla. The dioecious flowers of Sinilax are similarly complete, except by the abortion of one sex, but the calyx and corolla are single. FIG. 366. Flower of a Linum or Flax. 367. Andraecium and gyncecium ; the former of 5 perfect stamens, alternating with 5 rudiments of a second set. FIG. 368, 369. Dioecious flowers of Moonseed, Menispermum Canadense : 368, Stami- nate or male blossom; 369, Pistillate or female, but with six abortive stamens, before as many petals. FIG. 370. A catkin of staminate flowers of a Willow, Salix alba. 371. A single staminate flower detached and enlarged (the bract turned from the eye). 372. A pistil- late catkin of the same species. 373. A detached pistillate flower, magnified. 194 THE FLOWER. :;."»1. Combined with suppression of Perianth. This, which is found in most amentaceous or catkin-bearing trees, in some with partial suppression of perianth, is well illustrated ia Willows, the flowers of which are all achlamydeous and dioecious. (347.) The little scale (gland or nec- tary) at the inside of each blossom might be sup- posed to represent a perianth, reduced to a single piece ; but an extended comparison of forms refers it rather to the receptacle. Willow-blossoms (Fig. 370-:'.7-_) t are crowded in catkins, each one in the axil of a bract : the staminate flowers consist of a few stamens merely, in this species of only two, and the pistillate of a pistil merely. In Salix purpurea, the male flower seems to be a single stamen (Fig. 374) ; but it consists of two stamens, united into one body. Here extreme suppression is ac- companied with co- alescence of the existing members. :'..">.">. Still more simplilied flowers, but more dilh'cnlt to comprehend, arc those of Euphorbia, or Spurge. These are in fact monoeci- ous ; and the female flower is a pistil, the male is a stamen. The pistillate flower (of three carpels, their ovaries united into one three-lobed compound ovary) surmounts a slender peduncle which terminates each branch of the flowering plant. (Fig. 375.) From around the base of this pe- duncle rise other smaller and shorter peduncles, each from the axil of a slender bract, and surmounted by a single stamen, which represents a male (lower. (Fig. 376, 377.) This umbel-like I-'IC. :;7I. A separate staminate tl..\ver of Salix purpurea, with the stamens coa- lescent (mcmadelphous and syiiLrenesious). sn MS to appear like n single one. FK!. :!7r>. Flowering branch of Knpliorbia corollata. 370. Calyx-like involucre ilivideil lengthwise, showing the staminate (lowers around a pistillate flower (a). 377. A more magnified staminate Mower detaehed with its bract, a; its peduncle or pedicel b, surmounted by the solitary stamen, r. 378. Pistil in fruit, cut across, showing the three one-seeded carpels of which it is composed. 375 S78 SUPERPOSITION OF SUCCESSIVE PARTS. 195 flower-cluster is surrounded and at first enclosed by an involucre in the form of a cup, which imitates a calyx ; and the lobes of this cup (the free tips of the calyx-leaves) in the present species are bright white, so that the}- exactly imitate petals. Here, then, is a whole cluster of extremely simplified flowers, taking on the guise of and practical!}- behaving like a single flower, the invo- lucre serving as calyx and corolla ; the one-stamened male flowers collectively imitating the audroecium of a polyandrous blossom, and surrounding a female flower which might pass for the pistil of it. A series of related forms, from various parts of the worldT gives proof that this interpretation is the true one. 356. Suppression of both Androacium and Gfynoecium. This occurs in what are termed Neutral Flowers (347) , such as are conspicuous at the margin of the cymes of Hydrangea (Fig. 293) and of Viburnum lantanoides and Opulus, also at the margin of the head of flowers of Sunflower, Coreopsis (Fig. 287, 288), and the like. In these and most other instances, the perianth of which only the flower consists is much larger and more showy than in the accompanying perfect flowers : in fact, their whole utility to the plant, so far as known, is in this conspicuousness. No plant normally bears neutral flowers only ; but in cultivation all sometimes become so by monstrosity, as in the form of Vibur num Opulus called Snowball or Guelder Rose, also in "full double " roses, pinks, &c. Occasionally flowers become sterile and neutral by mere depauperation and abortion of perianth as well as of essential organs, as in certain Grasses ; but such are mostly vestiges of flowers rather than neutral blossoms. § 6. INTERRUPTION OF NORMAL ALTERNATION. 357. Anteposition or Superposition is the opposition of succes- sive (or apparently successive) whorls which normally alternate. This result is brought about in different ways, some of which are obvious, while of some the explanation is hypothetical. 358. In the first place, there are cases of seeming anteposi- tion, which are explained away on inspection. In a tulip, lily, and the like, there is a perianth of six leaves and a stamen be- fore each. The simple explanation is that the flower is not 6-merous, but 3-merous : there is a calyx of three sepals, colored and mostly shaped like the three petals, which alternate with these and are clearly anterior in the bud ; next, three stamens alternate with the petals or inner circle of the perianth ; then the three stamens of the inner circle, alternating with the preced- ing, necessarily are opposite the three petals, as the first three are 196 THE FLOWER. opposite the sepals. These organs altogether are in four whorls of three, not in two of six members ; and the pistil at the centre, of three combined members, is the lil'th and final whorl. 359. The Barberry family exhibits a similar seeming ante- position, which is more striking on account of a multiplication of the members of the perianth. The calyx is of six sepals in two circles, the corolla of six petals in two circles, the stamens equally six ; and so each petal has a stamen before and a sepal behind it. But. when properly viewed as a trimerous flower with double circles of sepals and petals as well as of stamens, all is symmetrical and normal. Menispermum in the related Moonseed family is in the same case, but the flower is trimerous, as seen in Fi<>\ .'$(>!> : in the male blossom this is obscured in the andne- ~ cium (Fig. 368) by a multiplication of the stamens.1 The same thing occurs in the perianth and bracts of certain ( 'lusiaceae, in which the members counted as in fours are superposed, and in some of which the double dimerous arrangement with apparent antcposiHon extends through the corolla ; while, in other closely related Mowers, the corolla changes to simply tetrainerous and to alternation with the preceding four sepals. This passes, in the same family and in the allied Ternstnemiaceae, into 360. Superposition by Spirals, as where live petals are ante- posed to five sepals, by an evident continuation of pentastichous phyllotaxy : and the stamen-clusters of Gordonia Lasianthus are probably in this way brought before the petals.2 The flower of Camellia is continuously on the spiral plan up to the gynce- cinm ; but upon one which, from the 1 tracts onward, rises from the \ to the I and £ order or higher, tin-owing the petals of the rosette in a full-double llower into numerous more or less conspicuous vertical ranks. 361. Anteposition in the Androeeiiim. It is in the andru-cium thai real anteposition is most common, and also most difficult to account for upon any one principle. Doubtless it comes to pass in more than one way. This condition is chiefly noticed when the stamens are definite in number, and mainly in isostemonous and diplostemonous flowers. (324.) 362. With Isoslemony. Vitis (Fig. 379-381 ). also Khamnus (Fig. 1 1 "). 1 16). and the whole Grape and Buckthorn families of 1 In Columbine (Aquilegia), multiplication of the stamens in successively alternating "nnerous whorls similarly brings the andnrrium into ten ranks; so, -when these stamens in double tl<>wers art- transformed into hollow-spurred petals, these are set one into another in ten vertical ranks. '-' Gen. Illustr. ii. t. 140. But the petals alternate with the sepals in the ordinary manner of the (lower, though their strong quincuncial imbrication suggests the spiral arrangement. SUPERPOSITION OF SUCCESSIVE PARTS. 197 which they are the types, afford familiar cases of a single circle of stamens placed before the petals. In Vitis, there are green nectariferous lobes or processes from 3^9 sso the receptacle, alternate with and inside the stamens : there is no good reason to suppose that they answer to a second row of stamens. All isostemonous Por- tulacaceae have the stamens before the petals ; and. when the stamens are fewer than the petals, those which exist occupy this position. Among the orders with gamopetalous corolla, such anteposition is universal in Plumbaginaceae, Primu- laeese, the related Myrsinaceae, and in X^vo most Sapotacea?, in the latter usually — •* with some complications. SS1 363. The earliest and the most obvious explanation of the anomaly is that of the suppression of an outer circle of stamens, and to this view recent morphologists are returning.1 Observa- tion supplies no vestige of proof of it in Rhamnacese and Vitaceae ; but, in the group of related orders to which the Primulaceae be- long, evidence is not wanting. For Samolus and Steironema both exhibit a series of rudimentary organs exactly in the place of the wanting circle of stamens, which may well be sterile fila- ments. In the allied order Sapotaceae, while Chrysophyllum has in these respects just the structure of Primulaceae, and Sideroxylon that of Samolus, Isouandra Gutta (the Gutta-percha plant) has a circle of well-formed stamens in place of the sterile rudiments of the preceding ; that is, alternate with the petals, 1 Eichler, Bliithendiagramme, passim, and in preface to Part II. xviii., relating chiefly to obdiplostemony. The principal opposing view is that of St. Hilaire, Duchartre, £c., maintaining that corolla and stamens here repre- sent one circle of organs doubled by median chorisis ; upon which see note under a following paragraph. According to that hypothecs, there is no androecial circle in such blossoms, or only vestiges of one, but the petals have supplied the deficiency by a supernumerary production of their own ! The more plausible hypothesis of Bratin, that of a suppressed interior circle of extra petals, would restore the alternation, and make the extant sta- mens the fourth floral circle, as does the adopted explanation. Braun's hypothesis, if it insists that an extra row of petals is wanting, supposes the suppression of that which very rarely exists ; but, if of stamens, then the supposed suppression is of that which is so generally present, or with indi- cations of presence, as properly to be accounted a part of the floral type. FIG. 379. Flower of the Grape Vine, casting its petals before expansion. 380. The same, without the petals: both show the glands of the disk distinctly, within the stamens. 381. Diagram of the flower. THE FLOWER. completing the symmetry of the blossom ami the normal alterna- tion of its members. This explanation of the anteposition of a single circle of stamens is the more readily received, because it well accords with the idea here adopted, that the andneeinni of a typical flower should consist of two circles of stamens. (•"•:.' 4.) The only serious objections to this explanation rise out of the diilieulty of applying it to analogous anteposition when both circles are present. 364. For Diplostemony^ the condition of two circles of sta- mens, each of the same number as the petals. i> al-o itself very commonly attended by anteposition. In normal or Direct Di/tlostemony, — that which answers to the floral type com- pletely,— the antisepalous stamens (324, note) are the outer and the antipetalous the inner series, and the carpels when isomerous alternate with the latter and oppose the sepals ; the alternation of whorls is therefore complete, as in the diagram, Fig. 382. Such stamens, however, may actually occupy a single line or coalesce into a tube, without derange- ment of the type. But it as commonly occurs that the antipetalous sta- mens are more or less exterior in insertion, and then the carpels, when isomerous, are alternate with the inner and anti- sepalous stamens, and therefore opposite the petals, as in the diagram. Fig. 383. This arrangement takes the name of Obdiplostemony. In it the normal alternation of successive whorls is interrupted, so as to produce anteposition, 365. With Olidiplostemony. This condition prevails, more or less evidently, in Krieaeea-. ( ieraniaeea', Xygophyllacea-. Kutaeejv, SaxilVagaeeie. ( Yassiilaee:e, ( )nagraecH', &c. (but in some of these with exceptions of direct diplostemonv) ; also, accompanied by a peculiar multiplication of members (380). in Malvacea-. Stercnliaee;e, and Tiliacea-. The explanation is dillicult. The hypotheses may be reduced to three, neither of which is quite satisfactory. There is. first, the hypothesis of St. Hilaire. ap- plied to this as to the preceding case (to Rhamnus, Vitis, &c.), that these exterior antipetalous stamens belong to the corolline whorl; in other words, that the petal and the stamen before it FIG. 382. I)i:iu'i:nn <>t i-uttern flower with direct diplostemony 383. Diagram of similar flower with ubdiplostemouy. Both from Eichler's Blutlu'iuliagrainme. SUPERPOSITION OF SUCCESSIVE PARTS. H>9 (whether aclnate to or free from it) answer to one leaf which has developed into two organs by a deduplication (372) taking place transversely. This makes the inner and antisepalous stamens the third floral circle or the only truly androecial one, and sym- metrically alternate with the petals on the one hand and the carpels on the other. The second hj'pothesis conceives that there is a whorl suppressed between these antipetalous stamens and the corolla : this, ideally restored, gives symmetric succession and alternation to all the succeeding whorls. The five glands in a Geranium-flower, alternate with and next succeeding the petals (Fig. 384), were plausibry supposed to represent this missing whorl, which according to Braun should be an inner corolla ; according to others rather a primary circle of stamens. The third is the recent hypothesis of Celakowsk}-, which Eichler adopts : this regards the antipetalous stamens as really the inner or second circle, and conceives that in the course of development it has become external by displacement. The difficulties of this hypothesis are, first to account for this dis- placement, and then for the anteposition of the carpels to the assumed inner stamens in the great majority of these cases.1 1 In the first part of the Bliithendiagramme, Eichler inclined to the first hypothesis, that of St. Hilaire (now very much abandoned on account of the feeble evidence that there is any such thing as transverse or median chorisis); in the second, he discards this in favor of Celakowsky's view (published in Regensburg Flora, 1875). As to members which are morpho- logically interior becoming exterior by outward displacement, Eichler cites the staminodia or sterile stamen-clusters of Parnassia (Fig. 400, 401), and the corresponding antipetalous stamens of Limnanthes, as clearly interior in the early flower-bud, but exterior at a later period ; states that the vascu- lar bundles which enter these stamens generally are either inner as respects those of the episepalous stamens or in line with them ; that in some cases (as in many Caryophyllacese) the real insertion of the stamens is that of direct diplostemony, while the upper part of their filaments and the anthers are external to the episepalous series; that in most families with obdiplos- temony examples of direct diplostemony occur, and still more cases with both stamineal circles inserted in the same line ; and that, as a rule, the episep- alous stamens are either later or not earlier formed than the epipetalous. As to the position of the carpels before antipetalous stamens and petals, Celakowsky suggests that this may result from the outward recession of those stamens affording more room there, while in the normal case the greater space is over the episepalous stamens. And, indeed, exceptions to the prevalent position are not uncommon both in direct diplostemony FIG. 384. Diagram (cross-section) of the flower of Geranium maculatum, exhibiting the relative position of parts, and the symmetrical alternation of circles, i. e. sepals, petals, greenish bodies called glands, antipetalous stnmons, untisepalous stamens, carpels. 200 THE FLOWER. 366. The case of stamens in a cluster before the petals is a complication of cither of the foregoing with a peculiar kind of multiplication, termed deduplication or chorisis. (372.) § 7. INCREASED NUMBER OF PARTS. 367. Augmentation in the number of floral members' is one of the commonest modifications of the type. It occurs in two ways : 1st. by an increased number of circle* or turns of spirals in the flower, which is Regular M/ilti/>lico that then' are transitions, a> above mentioned, between ohdiplostcinony and direct diploMemony. To Brann's theory that the glands behind the antiaepalous staniens in true Geraniaceie answer to suppressed phvlla, Kichler objects that these are present behind all ten stamens in <>xalidca>; also that all are wanting when the olliec of nectar-secretion, which they sub- serve, is undertaken by some other part of the flower, as by the calyx-spur in Pelargonium and Tropa>olum. The first objection is forcible: the second mixes morphological considerations with functional, and is inconclusive. Abortive organs, preserved for their utility as nectaries, might totally dis- appear when rendered useless by a different provision for the same function. INCREASED NUMBER OF PARTS. 201 is even more true of greatly multiplied stamens and pistils, as in Magnolia and Liriodendron, most Anonaceae, Ranunculus, Anemone, and the like. But in Aquilegia, where the number live is fixed in the perianth, the cyclic arrangement with alternation of whorls prevails throughout 369. The definite augmentation of calyx and corolla by the production of one additional whorl of each, and the seeming anteposition which comes of it when the androecium remains simply diplostemonous (in the manner of the Berberidaceae, Menispermaceae, &c., 359) has already been explained. 370. Similar increase to two whorls affecting the corolla only characterizes Anonacese, Magnoliaceos, Papaveraceae, and Fuma- riaceae. In all but the last order, this is accompanied by indefi- nitely multiplied stamens, and mostly by an increased number of carpels. In Fumariacese, which has dimerous flowers, there is a diminution by the suppression in most cases of half the normal androecium, and also an augmentation of the other half by chorisis. (372.) 371. Parapetalous Multiplication. Under this head may be described an anomalous arrangement of augmented stamens which prevails in the order Rosaceae, but is not peculiar to it.1 The simplest case, but a rare one, is seen in the 10-stamened vari- ety of some Hawthorns, as occasionally in Crataegus coccinea and Crus-galli. The ten are in one circle and in pairs, the pairs alternate with the petals. Some would say the pairs are before the petals ; but the space between two stamens before each petal is mostly rather wider than in the pair taken the other way. The next case in order, as in 15-starnened Hawthorns, and constantly in Nuttallia, adds to the above a simple interior circle of five stamens, one directly before the middle of each petal. Next, as in most Pomeae and many Potentilleae, there are twenty stamens, thus placed, but with an additional circle of five alter- nating with the preceding one. Next there are 25 in three circles, the second circle as well as the first having ten stamens ; and finally there are from 30 to 50, all probably in circles of ten each. There is little doubt that the circles develop in centri- petal order ; the inner successively the later.2 1 It was first clearly described by Dr. A. Dickson, in Trans. Bot. Soc. Edinb. viii. 468, and Seemann's Jour. Bot. iv. 473 (1860). He introduced the term, parapetalous, which is characteristic of it in its elementary form (254, note) : it is particularly illustrated by Eichler, in Bliithendiagramme, ii. 495-510. The former interprets it by chorisis, both median and collateral : the latter presents the facts and possible views, but declines to adopt either of them. 2 Accordingly, the whole is probably to be explained by some modifica- 202 THE FLOWER. 372. fhorisis or Dcduplication. Both these terms, and tin- ideas which they denote, originated with Dunal, but were first expounded by Moquin-Tandon.1 The first word is Greek for a separating <>r separation. The second is a translation of Dunal's French word dedoublement (literally undoubling). the ambiguity of which, and of the original presentation of the case, long retarded the right apprehension of the subject. Diremption has been suggested (by St. Ililaire) as a proper term. The mean- ing simply is, the division of that which is morphologically one organ into two or more (a division which is of course congenital), so that two or more organs occupy the position of one. As thus used, chorisis is restricted, or nearly so, to the homologues of leaves in the Mower, and mainly to stamens and carpels ; the division or splitting up of a petal or a sepal, when it occurs, being expressed in the phrases which arc applied to leaves. Yet, a compound leaf, especially one of the palmate type, is a good type of chorisis, the several blades of a compound leaf answering to the single blade of a simple leaf. Lt has been ob- jected against the terms chorisis and deduplication that they assume the division of that which has never been united; but so equally does the established terminology of foliage. A di- vided leaf has never been entire. 373. C'horisis is complete when the parts concerned are dis- tinct or separate to the very insertion, as in the stamen-clusters of Ilypericum. The foliar form of this would be represented by tion of the augmentation of circles. Dickson's hypothesis, that the two, three, or five stamens which are more or less in face of each petal are all deduplicationa of that petal, would come to be noticed under the next head, but it may be dismissed at once. Yet that the pairs in the outer circle represent each an antisepaloiis stamen, divided by chorisis (.-omeiimcs incompletely) and much separated, is not improbable. The other tillable explanation (which may be harmoni/ed with the last) is that the outer circle of stamens here rightly consists of ten members, respectively alternat- ing with the sepals and petals taken as a whole. This makes them para- pctalons, and at the same time brings them under 1 Int'meister's general law that new organs originate over intervals of those preceding, in this case over the ten perianth-intervals directly. It also accords with Hartog's elucidation of the accessory parts in the flower of Sapotacca- (in Tri men's Jour. Hot. IhTN). The inner circles are there sometimes ">-nicrous after the primitive type, sometimes iMmeroiis in regular alternation to the preceding circles. 1 Moquin-Tandon, I-'.-sai des De'donblemens, &c., Montpellicr, IS'JIi; Con- siilc'rations snr les Irrcgnlarites de la forolle, &e., in Ann. Sci. Nat. xxvii. 237, 1S:!-J; Teratologic Yegrtale, .'5:57 . Dunal, Kssai Mir les Vaccinie'es, IH'.I, cite, I by .Mnquin ( some pages printed, but never published); Conside- rations sur la Nature et les Rapports de quelques-nns des Organes de la Fleiir, 1S'2!>. The next botanist to develop it was St. Ililaire, Morphologic Ve'gctale, 1841. CHORISIS OR DEDUPLICATION. 203 such sessile pahnately compound leaves as those of some species of Aspalathus. It is incomplete when division does not extend to the base; as in Fig. 387, 393. Compare, as a proximate homologue of this, a petal of Mignonette, Fig. 385. But proper chorisis requires that the supernumerary organs should be developed like unto the original organ which is thus multiplied, or should complete their symmetiy, whatever it be. 374. St. Hilaire distinguished two kinds of deduplication ; viz., collateral when the members stand side by side, and pamllil when an organ becomes double or multiple antero-posteriorly. The latter, sometimes called vertical, and sometimes transverse, is better named median chorisis. The collateral is the origi- nal and typical chorisis. Most botanists incline to restrict the name to this, and to give some other explanation and name to the median form of augmentation. But some cases, such as those of Tilia and Sparmannia, are clearly of the same nature as the collateral, and ma^y be a disguised form of it ; there are others which may be explained in accordance with it ; and there^ are such transitions between some of these and coronal out- growths that the term chorisis is most conveniently made to comprise augmentation or doubling in either plane. Distinct anteposition, however, may be explained in other ways. (357.) 375. Typical or Collateral Chorisis, in which the members, together answering to one leaf, normally stand side by side, occurs in many families of plants, and in a variety of forms. A few are here presented. 376. Elodes Virginica (a common marsh plant of the Hypericuni family) , like most of its near relatives, has its calyx and corolla on the plan of five, its stamens and carpels on the plan of three, as is shown in the diagram, Fig. 38G. This makes a break in the symmetry between the corolla and the stamens; but all within is in regular alternation when the three stamens of each cluster are counted as one as their union at base into a plmluur (Fig. 387) may suggest. These phalanges alternate with the three carpels, and therefore stand where single stamens belong. The three conspicuous green projections, which in a general \v;iy FIG. 385. A petal of Mignonette (Reseda odorata), with many parted blade, enlarged. FIG. 386. Diagram of flower of Elodes Virginica, with three phalanges of stamens forming the inner circle, and three glands answering to the outer circle. 387. A de- tached phalanx of three stamens. as? 204 THE FLOWER. are called glands, alternate with the phalanges, and so are taken to represent the outer circle of stamens. The morphologist accordingly sees in the glands the homologues or representatives of the outer series of stamens, reduced to three by abortion, and in the three stamen-clusters only the three alternating stamens <>(' the inner series, trebled by chorisis, and this chorisis incom- plete, because it has not quite divided the filament into three. In Ilypericnni, the glands are completely suppressed, each pha- lanx is almost or quite divided into a cluster, either of about three stamens each, as in II. Sarothra, or of a few more (in II. nmtilum and H. Canadense) , or of an indefinite number, as in the common St. Johnsworts. Then in some other species (as in our II. pyramidatum) the carpels and the stamen-clusters rise to five, realizing complete pcntamerous symmetry, except that the almost numberless stamens all belong to the one inner circle. Morphologically, they are comparable to the leaflets of five (or in most species three) decompound and ses- sile or almost sessile leaves. The indefinitely numerous stamens of Ricinus are similarly increased from five by compound ramification. 377. Fumariaceoe, the Fumitory family, may furnish the next illus- tration. The flower is on the plan of two (dimerous) throughout. Taking Dicentra to show it. there is first a pair of small and scale-shaped sepals, not unlike the pair of bractlets on the FIG. 388. Dicentra ( 'urullai i:i ( Dutchman's Breeches), a scape in flower and a leaf, severed from tin- singular bnlb (formed of tin- enlarged bases of petioles). 389. Detached flower, of natural si/.e. showing also the pair of bractlets on the pedicel. 390. Same with parts displax ,-d, and :!!H. inner petals placed above. 392. Diagram of (lower of Dicentra <>r Adlnniia. lY.mi a section across the summit. 393. One of the phalanges of stamens of Adlumia; upper part only. CHORISIS OR DEDUPLICATION. 205 pedicel below (Fig. 389, 390), and normally alternate with them : alternate with these is a pair of large petals, deeply saccate or spurred below ; alternate with these, a pair of smaller petals with spoon-shaped tips which cohere at the apex (the corolla therefore of two circles as in the related Poppy family) ; alternate with these, two phalanges or united stamen-clusters, of three stamens each ; alternate with these is nothing, for the second set of stamens is wanting ; alternate with this vacancy is a pair of carpels wholly combined into a compound 2-merous pistil. The statement itself explains the morphology. The three sta- mens of each phalanx stand in the place of a stamen, and are the divisions of one. In Dicentra the members of the phalanx are almost separate ; in Adlnmia (Fig. 393) and Coiydalis the undivided filament reaches almost up to the anthers. The middle anther of the phalanx is normal, or two-celled ; the lateral anthers are one-celled, as if halved.1 1 Eichler adopts this interpretation (proposed in Gray, Gen. Illustr. i. 118), and applies it to the crucial instance of Hypecoum. In the flower of this Old World genus, there are four apparently simple and complete stamens, one before each petal : the simplest interpretation would be that which the facts appear to present, viz. that both dimerous circles of stamens are complete and normal. But Eichler — in view of the early development and the double vascular bundles of the stamens before the inner petals, and some occasional slight disjunction of their anther-cells — considers that the interior stamen-circle is wanting here, no less than in the other genera of the order ; that what here takes its place before each inner petal is a stamen composed of the adjacent lateral member of the phalanx, congeni- tally severed from the group to which it belongs and soldered into one fila- ment, bearing the two one-celled anthers so brought together as to imitate a normal two-celled anther. The organogeny of the blossom is thought to favor this hypothesis ; and it certainly favors the view here adopted of the composition of the three-membered phalanx of the family generally. If this interpretation of Hypecoum seems far-fetched, it is no more so than its exact counterpart, through which DeCandolle, Lindley, and others explain the case of the rest of the family. Starting with that genus as the simple type, they conceive that the stamen opposed to each inner petal is each severed into two, and that these half-stamens attached to the sides of the two intact stamens, thus producing the phalanges by coalescence. A good empirical conception of the formation, from a single leaf, of three stamens in Fumariaceze, or two in Crucifera, is afforded by the petals of Hypecoum, as illustrated by Eichler. The outer petals are slightly three- lobed from the apex ; the inner are deeply so and narrower. The mem- bers of the next circle in the family generally are just such three-lobed bodies, the tip of each lobe transformed into an anther. There is an ap- parent congruity in the production by the symmetrical middle lobe of a symmetrical two-celled anther, and of a one-celled anther by each unsym- metrical lateral lobe or stipule-like portion. A fuller derelopment of these sides of the leaf, and non-development of the middle portion (somewhat after the analogy of Lathyrus Aphaca, Fig. 219), with anther-formation, would convert the leaf into a pair of stamens. 206 THK FLOWER. 378. The obvious relationship of Cruciferse to Fumariaceae, their agreement in the rare ]icculiarity of having the two carpels side liy >ide in>tead <»f fore and aft (median), and the characteristic anomaly which the androecinm pre- sents (i.e. the tetradynamy) , would give reason to expect that its prob- lems might be solved by chorisis. «/ Indeed, the doctrine was applied to this, long before its application to the other order. Beginning at the centre (Fig. 395, . Diagram of such a tl<>u IT, with position of axis inarknl iiiiovc it. 396. Tetradynamons stamens and the pistil. 397. A common monstrosity of tin' same, two of (lie four inner stameua combined into a common 2-antlieriferous body. CHORISIS OR DEDUPLICATION. 207 single in Scnebiera and many species of Lepidium, in which the lateral or short stamens are at the same time abortive. 379. It is quite possible that chorisis may be extended to the corolla of the cruciferous flower, and reduce the whole to a symmetrical 2-merous plan, and to congruity in the perianth also with Fumariaceoe. The only obstacle is in the petals form- ing a whorl of four where all the rest is 2-merous, for the sepals are manifestly two decussating pairs. Now the median petals of Hypecoum are deeply 3-lobed. An abortion of their middle lobe would leave them almost two-parted : a little more would separate them ; then they would imitate the four cruciferous petals as in the diagram, Fig. 395. Applying this view to Crnciferse, the blossom in the two orders would accord in having a 2-merous three- whorled perianth, the first and third whorls median ; a as also in the dimerous audrcecium, the. first whorl of which is lateral. The difference is that in Fumariaceae the two members of the first whorl of stamens augment by chorisis into three, and the second is wanting, or is present only in Hypecourn ; while in Crucifera? the first whorl is simple (of the two short stamens) , and the second is doubled. In Furnariaceas only the first whorl of the perianth counts as calyx, and the corolla is of two whorls ; in Cruciferffi, the first and second whorls are calyx, the inner sepals answering to the outer petals of Fumariaceffi. 380. Chorisis along with anteposition of stamens is well seen in Tilia or Linden, at least in the American species. In these the indefinitely numerous stamens are in five clusters, one before each petal (Fig. 398, 399), and there is a petal-like body v x> alternating with the calyx-members as a whole ; the short stamens following as a 2-merous circle ; then the long stamens as a 4rmerous circle ; lastly the 2-merous gynoecium. G. Henslow (in Trans. Linn*. Soc. ser. 2, i. 195) would have the flower 4-merous by the suppression of the fifth members of a 5-merous type, and a further suppression of half of the remaining exterior stamen-circle, &c. Finally, there is the much better-maintained view that the cruciferous flower is 2-merous throughout, as explained in the following paragraph, 379. 1 This view was taken by Steinheil, :n Ann. Sci. Nat. ser. 2, 337 (1839), and is essentially reproduced by a Russian botanist, Meschajeff, in Bull. Soc. Imp. Nat. Mosc. 1872. FIG. 398. Diagram of the flower of Tilia Americana, the common American Lin- den or Basswood. FIG. 399. A detached stamen-cluster with its petal-like scale. 208 THE FLOWER. in each cluster with which the stamens cohere. The explanation by clmrisis is that each cluster, petal-like body included, is a multiplication of one stamen. The diagram (Fig. 308) accu- rately shows that most of the stamens originate from the outer side of the base of the petal-like portion : this is most naturally explained by median chorisis. The superposition of the clusters in i In- petals will take the same explanation as that of Rhamnus, Vitis, &c. (Fig. 363.) That the andrcccium is here composed of the inner circle' merely is partly confirmed by the alternation of the carpels with the clusters. According to Duchartre.1 the development of the andru'cimn in a Mallow indicates a similar structure ; for the whole united mass originates from five protu- berances, one before each forming petal and connected with it, this by collateral chorisis forming a cluster of stamens, and the five clusters coalescing as they develop into a tube of filaments, such as in Fig. 485. Now Hibiscus and its near relatives have a naked tip to the stamen-tube, ending usually in five teeth ; and Sidalcea, as is most strikingly shown in the C'alifornian S. diplosrypha, has two series of stamens, the outer (answering to those of Malva and its relatives) in five membrauaceous pha- langes, superposed to the petals ; the rather numerous inner series, more or less in phalanges, surmounts an interior filament- tube. Whence it is inferred that these, and the five teeth terminating the column in Hibiscus, represent the in- ner stamineal circle which is wanting in Malva, as it is in Tilia.2 381 . The case of Parnas- sia would be explained as analogous to that of Tilia, but with the stamen-clusters before the petals wholly sterile, and of fewer divisions, while an inner circle of five stamens 1 Comptes Rendus, 1844, & Ann. Sci. Nat. ser. .°,. iv. 123. Duchartre and others \vlio draw freely upon median chorisis to explain anteposition, and consider that congenital union proves it, take the phalanges in these cases, like the sin-le stamens in Vitis, to be an inner part of the petal itself. But this view appears to have had its day. 2 Gray, (ien. Illustr. ii. 44, 57, 7G-82. The position of the carpels before the petals in Pavmiia and Malvaviscus brings the former into symmetrical alternation with such an inner stamen-circle ; but it is not so in Hibiscus, which has the carpels before the sepals. KIO. 41111. A p. -tal .if Parnassia Curoliniana. with a triple staminodium before it. FIG. 4iil. Diagram of the flower of Paruassia Caroliniana. •tin. 41 IJ OUTGROWTHS. 209 alternate with the petals forms the effective anclroecinm. For the scale-like body before each petal, and even slightly adnate to its base (in P. Caroliniana about 3-parted, as in Fig. 400, but in P. palustris a thin scale, fringed with more numerous gland- tipped filaments), is plainly outside the stamens in the full-grown flower-bud. But Eichler and Drude have found that it is inside in the early bud.1 Wherefore, if these stamen-like bodies really represent a circle of the androecium, it must be the inner one ; and that is the more probable view. 382. Multiplication by chorisis in the gynoecium is not common ; but there are well marked in- stances of it in all degrees. In Drosera, the styles and stigmas are doubled (Fig. 402) ; in Malvaceae, the same thing takes place in Pavonia and its allies ; while in Malope and two other genera of the same order the few normal carpels are multiplied, evidently by chorisis, into an indefinite number of wholly distinct ones. § 8. OUTGROWTHS. 383. Proper chorisis is the congenital multiplication of one organ into two or more of the same nature and office ; or at least into two or more organs, even if dissimilar, as in the American Lindens, in which one member of the cluster is a kind of petal. Between this and the production by an organ of ap- pendages, or outgrowths of little or no morphological signifi- cation, there are many gradations ; as also between these and mere cellular outgrowths from the surface, even down to bristles and hairs. The latter, in all their variety and modifica- tions, are properly outgrowths of the epidermis only, and there- fore consist of extended cells, single or combined, unaccompanied by vascular or woody tissue. To them has been given the general name of Trichomes (Trichoma, pi. trichomata) , that is structures of which hairs are the type. The}' may occur upon the surface of any organ whatever. Their morphology is the morphology of cells rather than of organs. They will therefore be most convenient!}' illustrated under Vegetable Anatomy as 1 Eichler in Fl. Brasil., Sauvagesiacae, & Bliithend. ii. 424 ; Drude in Lin- naea, xxxix. 239. Eichler refers to this as a confirmation of Celakowsky's explanation of obdiplostemony by posterior displacement. (3G5.) FIG. 402. Pistil of Drosera filiformis with tricarpellary ovary (transversely divided), and six styles, i. e. three, and each two-parted. 210 THE FLOWER. their structure, and in the Glossary as respects ter- minology. 384. But into some bristles, such as those of Drosera. a sub- jacent stratum of tissue enters, including one or more ducts or even some woody tissue. Prickles are of this class; and from the most slender, which pass into bristles, there are all grada- tions of stoutness and induration. Such outgrowths ma v even be formed in most regular order, as the prickles on the calyx-tube of Agrimonia and scales on the acorn-cup of Oaks, and yet have no morphological importance. On the other hand, true represen- tatives of leaf or stem may, by abortion and depauperation, be reduced to the structure as well as the appearance of trie-homes. Examples of this are familiar in the pappus (answering to limb of the calyx) of many Composite, and in the bristles which answer to perianth in many Cj'peraceae. The scarious stipules of Paronychia and of Potamogeton, the ligule of Grasses, and even the corolla in Plantago, arc equally reduced to mere cellular tissue. So that the structural difference between trichomes and outgrowths l is not at all absolute, and the morphological distinc- tion must rest upon other ground than anatomical structure. 385. Among the corolline outgrowths most akin to chorisis is the Crown (Corona) of Silene and allied Caryophyllaceae, at the junction of the claw with the blade of the petals (Fig. 403), the analogy and probable homology of which to the ligule of Grasses (Fig. 150) is evident ; also the many-rayed fila- mentous crown of Passion-flowers (Fig. 404), which consists of two or more- series of such outgrowths. In Sapindus and some other Sapindace:e. these lignlar outgrowths or internal appendages are more like a doubling of the petal: as also in Erythroxylum, where they 1 This is the best English name for the Kmin/nizoi of the Germans, the K/>i!>/I. is a term properly applied to a short and comparatively broad portion of receptacle on which the gyixrcium rests, as in Hue and Orange (Fig. Ill), lloundstoiigue, Sage, &c. This may extend up between the car- pels and pass into, or the upper part become a CARPOPHORE, a name properly applied to a portion of receptacle which is prolonged between the carpels as a central axis, as in Geranium (Fig. 411) and many Umbellifene, Fig. 412. FIG. 408. Section of a flower of Silene Pennsylvania, showing the stipe or antbophore. FIG. 409. Flower of Gynandropsis, with floral circles separated on the elongated receptacle. FOKMS OF THE RECEPTACLE. 213 392. Instead of forming a stalk, the elongation ma}1 be continued between the carpels in the form of a slender axis, as in Gera- nium (Fig. 410, 411), and in the carpophore of the fruit of Umbelliferse, Fig. 412. In Geranium, this prolongation of receptacle 410 413 414 extends far above the ovaries as a beak, to which the styles are actuate for most of their length. 393. In Nelumbium (Fig. 413), the gynophore, or portion of receptacle above the stamens, is enlarged into a singular broadly top-shaped body, with a flat summit, in which the pistils (a dozen or more isolated carpels) are separately immersed. 394. A Disk is a part of the receptacle, or a development of it, enlarged under or around the pistil. When under it or around its base and free from the calyx, the disk is hypogynous, as in Orange, Fig. 414. Here it is a kind of gynobase. When adher- ent to or lining the base of the calyx, it is perigynoits, as in 4I5 Rhamnus (Fig. 415,416) and Cherry (Fig. 337) : when carried by complete adnation up to the summit of the ovar}T, it is epigy- nous, as in Cornus, in Umbelliferae, &c. Not rarely it divides into lobes, as in Vitis (Fig. 379, 380), in Periwinkle and most Apocynaceous plants, and in Cruciferse. These are termed (/lands of the disk, and indeed are commonly glandular or nectariferous. FIG. 410. Gynoeciurn of Geranium maculatum. 411. The same with fruit mature, the five ovaries or cells and the lower part of their styles separated and recurving away from the prolongation of the axis or receptacle, to which they were at flowering-time firmly attached. FIG. 412. Mature fruit, of Osmorrhiza, the two carpels splitting away below from the filiform prolongation of the receptacle, or carpophore. FIG. 413. The top-shaped receptacle of Nelumhium, with the pistils, immersed in hollows of its upper face. FIG. 415. Flower of a Rhamnus or Buckthorn, and 416, section of the same, show- ing a thickened perigynous disk. 214 THE FLOWER. It is not possible by any direct demonstration to distinguish be- tween such productions of the receptacle, which are classed as belonging to the axis, and suppressed or undeveloped phyllous organs, such as stamens, which glands of the disk may some- times represent. 39o. Hypanthium. Inspection of Fig. 415, 416, and 337, and comparison with Fig. 339, will suggest an explanation differ- ent from that which is generally adopted. Instead of regarding the calyx as beginning on a level with the base of the ovary, and the cup as lined, more or less thickly, by an expansion of the receptacle (the perigynous disk), the calyx may be understood to begin where this and the ovary become free from each other. I'liderthat view, the receptacle, instead of convex or protu berant, is here con- cave, has grown up 419 around the ovary, which, however, is free from the cup in the earlier cited figures, but immersed in it in Fig. 339 and the like. A comparison with a rose-hip, an apple, and a pear much strengthens this interpre- tation, which is rather largely adopted at this day. at least theoretically. It was perhaps lirst proposed by Link, who intro- duced the appropriate name of HYI-ANTIUM. A hypanthium or hypanthiol receptacle is, as the name betokens, a llower-axis or receptacle developed mainly under the calyx. The name is a good one, in any case ; and such structures as those of C'alycan- thus (Fig. 417-419), a rose, a pear (the lower part of which is evidently an enlargement of peduncle), and of Cactus-Mowers (Fig. •"• 17). although (mite compatible with the theory of actuation, are more simply explained by it.1 1 But, whether the cases are well distinguishable or not, it l>y no means follows that the receptacle plays such a part in all instances of perigyny and of inferior or partly inferior ovary. Such a view is attended by more diffi- culties than the other. Unless the mediation of an invisible receptacle must KKl. 417. Flowering branch of Calycan thus. 418. Vertical section of the urn-shaped receptacle, the imbricated bracts or sepals on its surface cut away. 419. Mature fructiferous receptacle entire, showing some scars from which the bracts have fallen. ADAPTATIONS TO FERTILIZATION. 215 SECTION IV. CERTAIN ADAPTATIONS OF THE FLOWER TO THE ACT OF FERTILIZATION. § 1. IN GENERAL. 396. The introduction into morphological botany of the con- siderations now to be mentioned should have dated from the year 1793, in which Christian Conrad Sprengel published his curious treatise on the structure of flowers in special reference to insect aid in their fertilization. For this book, which was wholly neglected and overlooked for more than sixty years, con- tains along with some fanciful ideas the germs of the present doctrine and many excellent illustrations of it.1 The interest in the doctrine now prevalent is witnessed by a copious special literature, beginning with the publication, in 1862, of Darwin's book on the fertilization of Orchids by the aid of insects.'2 be invoked whenever there is a junction of two dissimilar organs, the petals and stamens of a Lythrum or a Cuphea are united with the calyx itself, instead of calyx beginning at the top of a long and simple tube. And if three or more of the floral whorls may be congenitally united, why not these also with the remaining one 1 Van Tieghem, in his Anatomie Compare'e de la Fleur, maintains wholly the old view, founding it upon anatomical struc- ture and his ability to trace down to the base of the ovary the distinct vascular bundles of the several involved organs. 1 C. C. Sprengel, Das entdeckte Geheimniss der Natur im Ban und der Befruchtung der Blumen, Berlin, 1793. Even earlier, Koelreuter (Vorlaufige Nachricht, etc., 1761-1766) recognized the necessity of insect-aid to various blossoms, and described some special contrivances for the purpose. 2 Charles Darwin, On the Various Contrivances by which British and Foreign Orchids are fertilized by Insects, and on the Good Effects of Inter- crossing, London, 1862. Ed. 2, 1877. This last contains a list of the papers and books which bear upon the subject, published since 1862. Other leading works and papers on the subject are, exclusive of the other volumes and papers of Darwin, more or less referred to hereafter. Treviranus, Ueber Dichogamie, &c., in Bot. Zeitung, xxi. 1863. Hugo von Mohl, Einige Beobachtungen iiber dimorphe Bliithen, Bot. Zeitung, xxi. 1863. Delpino, Pensieri sulla Biologia Vegetale, &c., 1867. Relazione sull' Apparecchio della Fecondazione nelle Asclepiadie, &c., 1867. Ulteriore Osservazioni sulla Dichogamia, &c., 1868-69, 1870, and later papers. Axell, Om anordningarna for de Fanerogama Vaxternas Befruchtung, Stockholm, 1869. Hildebrand, Die Geschlechter-Vertheilung bei den Pflanzen, 1867, and other papers. Hermann Miiller, Die Befruchtung der Blumen durch Insekten, 1873, and papers in " Nature " and elsewhere. " Flowers and their Unbidden Guests," an English translation of a work by Professor Kerner, which describes arrangements in blossoms for exclud- ing unwelcome guests, has not yet reached us. It introduces the new terms Autogamy and Allogamy, defined on the following page ; the latter compre- 21G THE FLOWER. • 397. The subject, here considered as a part of morphology, must be fully treated, as regards acts and processes, under ).h\ Bi- ology. Every thing in the flower is in relation to fertilization and fructification, directly or indirectly. This section is con- cerned with those adaptations of structure by means of which agents external to the blossom are brought into service for its fertilization. 39 are either dissected into plumes, as in most Grasses, or beset with copious hairs on which pollen is caught. One physiological adaptation, very common in the fol- lowing class, is not unknown among hermaphrodite wind-fertiliz- alile flowers, where it is important for securing intercrossing, viz. Dichogamy. It is best seen in the common species of Plantago or Plantain, and is described below. (408.) 404. Insect-fort Hi/able or enlomophiloits flowers are correlated with showy coloration (including white, which is most showy at dusk), odor, or secretion of nectar, often by all three modes of attraction to insects combined. Sonic insects, moreover, visit flowers for their pollen, a highly nutritions article, and ordina- rily produced in such abundance that much may be spared. The showiness of corolla or other floral envelopes is an attractive. adaptation to fertilization, enabling blossoms to be discerned at a distance ; nor do we know that fragrance or other scent or that nectar subserves any other uses to the flower than that of alluring insects. Adaptations in the pollen of such blossoms for transportation by insects are various. Commonly the grains are slightly moist or glutinous, or roughish, or studded with projections, or strung with threads (as in (Enothera). so as not to be readily dispersed in the air. but to have some slight coherence as well as capability of adhering to the head. Mini's, or bodies of insects, especially to their rough surfaces; and in two families (( Mvhidacese, Asclepiadacene) the pollen is com- bined in masses and with special adaptations for being trans- ported en masse. (421.) With this the stigma is usually correlated, by roughness, moisture, or glutinosity.1 405. Adaptations of the flower itself in reference to insect visitation are wonderfully various; and most of these are found upon investigation to favor, or often to necessitate, intercross- ing. In diieeioiis (lowers, this is necessitated by the separation; in moiiiceiotis and polygamous flowers, of various kinds and 1 Tlm< nearly rvt-ry Orchid i/cnns lint one has a persistently glutinous stigma ; in tin- exceptional one, Cypripc'dium, it is moist and minutely rough- ened, in correlation with the loosely granular or pultaceous pollen which it is to receive. ADAPTATIONS FOR INTERCROSSING. 219 degrees of separation, pollen is very commonly borne from plant to plant ; in hermaphrodite flowers only are more special arrangements needed to secure intercrossing or a certain measure of it, and in these such arrangements abound. 406. Irregularity is one of the commonest modifications of the flower (326, 337) : it is never conspicuous except in blossoms visited by insects and generally fertilized by their aid ; and it finds rational explanation on the score of utility in this regard.1 407. Dichogamy, a term introduced b}' C. C. Sprengel, who first noticed and described it, is one of the most usual and effect- ual (rather physiological than morphological) adaptations for the promotion of intercrossing between hermaphrodite flowers. It means that such intercrossing is brought to pass by a difference in the time of maturity of anthers and stigma ; this rendering dichogamous blossoms practically the same as dioecious or mon- oecious in respect to fertilization, while there is the economical gain that all the flowers are fertile. According to whether the anthers or the stigmas are precocious, dichogamous flowers are Proterandrous (or Protandrous} , when the anthers mature and discharge their pollen before the stigma of that blossom is recep- tive of pollen ; Proterof/ynoiis (or Protogynous), when the stigmas are in receptive condition before the anthers have matured their pollen. Synanthesis* the maturing of the two sexes simultaneously or nearlv so, is however made to secure the same result through O special arrangements. 408. Proterogyny. The Plantains, such as Plantago major and P. lanceolata, are familiar instances of this in a wind-fertilized genus with hermaphrodite flowers. The anthesis proceeds from base to apex of the spike in regular order, and rather slowly. While the anthers are still in the unopened corolla and on short filaments, the long and slender hairy stigma projects from the tip and is receiving pollen blown to it from neighboring plants or 1 This did not escape the attention of Sprengel in the last century, and along with it the fact that strictly terminal and also vertical flowers, whether erect or suspended, are seldom irregular, while comparative!}- horizontal or obliquely set flowers more commonly are so. The irregularity is in refer- ence to a landing place for the visiting insect, or also to storage of or accessi- bility to nectar, &c. Darwin (Forms of Flowers, 147) remarks that he does not know of a single instance of an irregular flower which is wind-fertilized. 2 Synacmy is the term proposed by A. W. Bennett, in Journal of Botany, viii. (1870), 316, with its opposite, Heteracmy, for proterandry and proterogyny. The latter names, in their shorter form (protandry and protogyny), appear to have originated with Hildebrand, 1867. •2-20 THE FLOWER. spikes : a day or two afterwards, the corolla opens, the filaments greatly lengthen, and the lour anthers now pendent from them give their light pollen to the wind ; but the stigmas of that flower and of all below it on that spike are withered or past receiving pollen. Among Grasses, Anthoxanthum is in the same ca^e. The arrangement is somewhat similar to the Plantain in Amor- pha, which is fertilized by insects, the simple stigma projecting beyond the corolla in bud, while the anthers are still immature and enclosed. Scrophularia is a good instance of proterogyny in flowers fertilized by bees. The flower is irregular (Fig. 420-422), and is approached from the front, the spreading lower lobe being the landing place. Fig. 420 represents a freshly opened blossom; and Fig. 421, a section of it. Only the style tipped with the stigma is in view, leaning over the landing place ; the still closed anthers are ensconced below. The next day or a little later all is as in Fig. 422. The style, now flabby, has fallen upon the front lobe, its stigma dry and no longer receptive : the now-opening anthers are brought upward and forward to the position which the stigma occupied before. A honey-bee, taking nectar from the bottom of the corolla, will be dusted with pollen from the later flower, and on passing to one in the earlier state will deposit some of it on its fresh stigma. Self- fertilization here can hardly ever take place, and only through some disturb- ance of the natural course. 40!>. Proterandry. The process is the reverse, and is at- tended with much more extended movements in Clerodendron Thompsoniffi, a Verbenaceous tropical African climber now com- mon in conservatories. The adaptations in this flower (which we indicated long ago) are exquisite. The crimson corolla and bright white calyx in combination are very conspicuous. The long filiform filaments and style, upwardly enrolled in the FIG. 420, 421. Early opened flower of Scrophularia nodosa, and a longitudinal section. 422. Flower a day or two later. ADAPTATIONS FOR INTERCROSSING. 221 bud, straighten and project when the corolla opens : the stamens remain straight, but the style proceeds to curve downward and backward, as in Fig. 423. The anthers are now discharging pollen : the stigmas are immature and closed. Fig. 424 repre- sents the flower on the second day, the anthers effete, and the filaments recurved and rolled up spirally ; while the style has taken the position of the filaments, and the two stigmas now separated and receptive are in the very position of the anthers the previous day. The entrance by which the proboscis of a butterfly may reach the nectar at bottom is at the upper side of the orifice. The flower cannot self-fertilize. A good-sized insect flying from blossom to blossom, and plant to plant, must transport pollen from the one to the stigma of the other. 410. Proterandry abounds among common flowers. It is conspicuous in Gentians and in nearly all that family. But, while in Gentians the short style is immovable and erect, in Sabbatia it is thrown strong!}1 to one side, out of the wa}T of and far below the stamens, the branches closed and often twisted, so that the stigma is quite inaccessible until the stamens have shed their pollen : then the style becomes erect, untwists, its two flat branches separate, and expose the stigmatic surface of their inner face in the place which the anthers occupied. In Sabbatia angularis, Lester F. Ward 1 observed that the anthers of freshly 1 In Meehan's Gardeners' Monthly, September, 1878, 278. FIG. 423. Flower of Clerodendron Thompsoniae, first day; 424, second day. 222 THE FLOWER. opened blossoms are all thrown to one side almost as strongly as the style is thrown in the opposite direction. One of our common Fireweeds, Kpilobium angustifolium or F. spicatum, as it is variously called, which is common all round the northern hemisphere, is similar to Sabbatia in behavior. In the freshly opened flower, while the anthers are in good condition and are 425 426 giving their pollen to bees, the still immature style is strongly curved downward and backward, as in Fig. 425. Two or three days later, when the pollen is mostly shed, the style straightens, lengthens to its full dimensions, and spreads its four stigmas over the line of the axis of the blossom (Fig. 426), in the verv position to be pollinated by a bee coming from an earlier flower. 411. In the following instances of proterandry. the- style is made the instrument of distributing the pollen which it is not itself to use. The anthers of a Cam- panula discharge all their pollen in the unopened bud, and it is nearly all deposited on the style which 11 icy surround, the upper part of which is clothed with a coat of hairs for holding the pollen. (Fig. 427.) In the open flower, the stamens are found to be empty and withered, as in Fig. -12.s. These flowers are visited by bees and other insects for Hie pollen. While this is going on, and while the pollen is fresh and plentiful, no stigma is apparent. Later, the top of the style opens into three (in some species five) short and spreading branches, the inner faces of which are the stigmas. Although FIG. 425, 426. Flowers of Kpiloliium an-jiistifolium or spicatum; iu the first, freshly expanded : in tin- second, a few days older. Fl(}. 4'J7. Vertical section of an unopened flower of Campanula rapunculoides : the broad white lines are sections of two anthers. 428. Same of an older flower. ADAPTATIONS FOR INTERCROSSING. 223 so close at hand, little if any of the pollen of that flower can reach the stigmas. These actually get fertilized by pollen brought by bees, which come loaded with it from other flowers and other plants. Symphyandra differs from a true Campanula chiefly in the continued cohesion of the five anthers into a tube around the st}Tle. (Fig. 429, 430.) The pollen is discharged on and held by the hairy upper portion of the style. Soon after, the corolla expands, the lower part of the style lengthens, and carries the pollen-loaded part out of and above the anther-tube, as in Fig. 430 ; lastly, the three connivent tips of the style diverge and expose the stigmas to pollen mainly brought by bees from other flowers. By a slight further modification in Lobelia and in Composite, pollen is pushed out of the anther-tube by the tip of the style as it lengthens, or by the very back of the two stigmas, the faces of which, afterwards exposed, are not to receive this, but other pollen, though it may at times receive some of its own. The arrangement in Composite is here illustrated from Leptosyne maritima (Fig. 431-435), a showy plant of Southern California, now not very rare in cultivation. The large flowers around the FIG. 429. Stamens and pistil of a young, and 430, same from an old flower of Symphyandra pendula. FIG. 431. Head of flowers of Leptosyne maritima, of tbe natural size. THE FLOWER. margin (ray-flowers, with ligulatc corolla), one of which is sepa- rately shown in Fig. 432, are pistillate only : the enlarged and extended open part of the corolla (bright yellow in color) serves for attrac- tion, the circle of rays gives the appearance as of a single large flower. The flowers of the disk or whole central part are hermaphrodite, and with narrow tubular corollas. from the orifice of which projects the greater part of the tube of five co- alesccnt anthers. The pollen is early discharged into the interior of this tube. The style, with somewhat enlarged and brush-like tip, at tirst reaches only to the bottom of the anther-tube: it slowly lengthens, pushes the pollen before it out of the tube ( Fig. 433) and into the way of insects of various kind, which. travelling over the surface, con- vey it to older flowers of the same 432 433 434 435 head and of other plants. The style, elongating yet more, raises some of the pollen still higher (as in Fig. 434) : and at length its two branches separate and diverge (Fig. !.">'>). exposing to other pollen the stigmatic receptive surface which until now was un- approachable. 412. In Farnassia. which has sessile stigmas, their receptive surface is actually not formed until the anthers become elfete ; FIG. 4:!2 A ligulatc female flower of the same, anil a central hermaphrodite flower. 433. Upper part of tin- latter, more enlarged, tlio tube of anthers projecting from the corolla, and the pollen projecting from apex of the aiitlier-ttilie. being pushed tip by the lengthening of the style beneath. 4:vt. This style now projecting, and some pollen still resting on its tip 4:r>. Tip of same style (more advanced and magnified); the two branches spreading, still carrying some pollen on the apex of each arm or branch, by the divergence now exposing the stigmatic inner faces. ADAPTATIONS FOB INTERCROSSING. 225 and, as the plants or stems are single-flowered, they are function- al^- dioecious while structurally hermaphrodite. 413. The adaptations for hermaphrodite intercrossing with synanthesis (407), i. e. where there is no essential difference of time in the maturing of anthers and stigma, are manifold. They ma}' be classed into those without and those with dimor- phism of stamens and pistils, or, in other words, those with Homogenous and those with Heteroyonous flowers.1 414. The cases without dimorphism are the most various, certain families having special types ; and are of all degrees, from those that require intercrossing to those that merely favor or permit it. For the present purpose, having only morphology in view, it suffices to bring to view two or three cases or types of 415. Particular Adaptations in hermaphrodite blossoms, not involving either dichogamy or dimorphism. These are exceed- ingly various ; but they may be distinguished into two general kinds, namely: 1, where loose and powdery pollen is transported from blossom to blossom in separate grains, and 2, where pollen-masses or the whole contents of anthers are bodily so transported. 416. Papilionaceous flowers (such as pea-blossoms, 338) --having ten stamens enclosed with a single pis- til in the keel of the corolla, their anthers in close proximity to the stigma — were naturally supposed to be self-fertilizing ; and so they sometimes are, yet with marked adaptations for intercrossing. None are less so than those of 1 Terms proposed in Amor. Jour. Sci. ser. 3, xiii. 82, and in Amer. Naturalist, January, 1877. Dimorphism in flowers may affect the perianth only, and not the 701/7? or essential organs ; or there may be two kinds of flowers as respects these also, but with no reciprocal relations, as in cleisto- gamous dimorphism (534) ; or of two kinds essentially alike except in stamens and pistil, and these reciprocally adapted to each other, which is heterogonous dimorphism, or, when of three kinds, trimorphism, FIG. 436. Flower of Wistaria Sinensis natural size. 437. Same enlarged, with standard, wings, and half the keel removed. 438. Same with the keel depressed, as it is when a bee alights on this its usual landing place, the cluster of anthers and stigma thus brought up against the bee's abdomen. 439. Style and stigma, with part of the ovary, more magnified, a fringe of flue bristles around the stigma. 15 226 THE KLCAVEJK. 442 Wistaria (Fig. 436-439). in which the light fringe of stiff hairs around the stigma (shown in Fig. 43'J) would not prevent pollen of surrounding anthers from falling upon it. Yet when a liee alights upon the ked, with head toward the base of tin- (lower, and proboscis is inserted for nectar between the foot of the standard and the keel, the latter is depressed b}r the weight, so that the ab- domen of the insect is brought against the ten anthers and the stig- ma, becoming thereby smeared with pollen. some of which when other blossoms are vis- ited cannot fail to be applied to their stigmas. The very similar flower of Locust (Robinia). like that of the Pea, adds an adapta- tion in favor of intercrossing. The style for some length below the stigma is covered with a short beard of hairs, as is seen in Fig. 442. The anthers open early and dis- charge their pollen, which mainly lodges on this beard (Fig. 443), in a manner which may thus far be likened to the case of Campanula. (411.) The wings and the keel are yoked together, and are together depressed by the weight of an alighting bee. This docs not bring out the anthers as in Wistaria, but these remain until effete within the sac, while the stigma and the pollen-laden part of the style (Fig. 441) are projected against the bee's abdomen, which, by the oblique movement, is first touched by the stigma and next brushed over with pollen by the style below. So that, in visiting a succession of blossoms, some pollen of one flower is transferred to the body of the bee. and thence to the stigma of the next (lower, which (lower immediately gives to the same spot some of its pollen, to be transferred to (he next flower's stigma, and so on. 417. Two special modifications of the papilionaceous type FI«. 440. Flower of Koi.inia hiapida, the standard and frings removed. 441. Same, ;m depressed liy Ilir Wright ol a lice, i .IIIMIIL,' tin- stigma anil pollen-laden tip of the Style to protrude. 442. Kn larked section of same in the bud, leaving "lie keel-petal, half the KiaineiiH, and the pistil in view. 443. Style and stigma at a later period, the beard loaded with pollen; more magnified. ADAPTATIONS FOK INTERCROSSING. need particular mention. One of them, the Bean-blossom, is well known to botanists ; the other not so. The peculiarity in the common Beau, Phaseolus vulgaris, and its nearest relatives, is that the keel, enclosing the stamens and pistil, is prolonged into a narrow snout which is spirally coiled (as in Fig. 444-446) ; that the stigma is oblique on the tip of the style, and the beard on the style is mainly on the same side that the stigma is : the wing-petals stand forward and turn downward, forming a con- venient landing place for bees. As in the Locust-blossom, the anthers earl}* discharge their pollen, much of which adheres 448 lightly to the beard of the style. In the untouched flower, all from first to last is concealed in the coiled keel. Press down the wing-petals, and first the stigma and then the pollen-laden tip of the style projects from the orifice : remove the pressure, and they withdraw within. When this pressure is made by a bee, resting on the wing-petals while searching for nectar within the base of the blossom between the keel and the standard, the same movement occurs : the stigma first, and then the pollen on the style, strikes against a certain portion of the front or side of the bee's body, and the repetition of this operation causes the fertilization of each blossom by other than its own pollen. A slighter pressure or lighter movement of the wing-petals suffices FIG. 444. Flower of Garden Bean, Phaseolus vulgaris. 445 Same with wing- petals pressed down and tip of style projecting from the orifice of the keel. 446. Same as 444 enlarged, and standard and wings removed. 447. Upper part of keel, in the condition of 445, enlarged, showing plainly the projecting style. 448 Section of the keel, enlarged, showing the style within before the anthers open: stamens for sake of clearness not delineated. 449. Pistil detached from an older flower; the brush loaded with pollen. 228 THE FLOWER. to jostle some of the pollen down upon its own stigma, so that sell-fertilization is not uncommon. 418. Apios tuberosa. a near relative of Phaseolus, exhibits a different and equally curious modification of the same parts. The- wing-petals for landing place arc .similar: the standard is pro- portionally large, firm in texture, and shell-shaped or concave, with a small boss at the tip as seen from In-hind, or a shallow sac as seen 45! from the front : the keel is narrow and sickle-shaped ; it arches across the front of the flower, and the blunt apex rests in the notch or shallow sac of the tip of the 452 standard. (Fig. 450, 452, 453.) So it remains if untouched until the blossom withers : no self-fertilization has ever been observed, and none ordinarily oeeiirs. The anthers are assembled close around the stigma, but a little short of it (Fig. 452) ; the pollen is not early nor copiously shed in the enclosure: the small terminal stigma is at first covered with a pulpy secretion, which at length collects into a soft ring around its base over or through which no pollen passes. But when the keel is liber- ated by lifting from underneath, it curves promptly into the shape shown in Fig. FTO. 450. Flower of Apios tuherosa. unvisited. 4fd. Same after visitation, the ke.-l dislodged from the rctainim: notch. :ind more incurved; tin- tip of the style pro- truded :uid thrust forward, follow. •.] l.y the anthers Kl<; •(. -.•_'. Enlarged vertical section of flower-bad of Aploatuberosa. -l"3 A flower with half the standard cut away, to show the hlnnt apex of the keel resting in the notch. 4.-.I. Diagram of (lower, with half of the standard cut away, toshow what takes pla.-e when the apex of the keel is liberated. The figures (also those from 4'-'3 to the present!, and the flrel account of the adaptations of Apios, were published in the Amer- ican Agriculturist in 1876. ADAPTATIONS FOR INTERCROSSING. 229 451, or better in Fig. 454, where the dotted lines indicate its original position ; and first the end of the style, tipped with its stigma, is pushed forward, and then the anthers come into view. The flowers are visited by humble-bees, and sometimes by honej'-bees. In searching for nectar at the base of the flower, they probably push forward into the space under the arching keel, and by slightly elevating dislodge its apex ; when first the stigma and then the anthers are brought against some portion of the insect's bodj', and against the same portion in succeeding blossoms, thus effecting cross-fertilization. This rationally ex- plains a remarkable adaptation, which seems to be not otherwise intelligible. 419. Special Adaptations. Two of these, each peculiar to the genus, may here be referred to. In Kalmia-blossoms (Fig. 455 456 457 455-458), the anthers discharge the pollen through a small orifice at the apex of each cell, in this respect agreeing with Rhododendrons and their other relatives ; but none of them utilize this family peculiarity in the manner of Kalmia. In the flower-bud, each of the ten anthers is lodged in a small cavity or pocket (external!}- a boss) of the corolla, in a way analo- gous to that in which the keel of Apios is lodged in the tip of the standard (418) : the expansion of the border of the corolla in anthesis curves the fila- ments outward and backward ; and when the bowed 453 stamens are liberated by rough jostling they fly up elastically, and the pollen is projected from the two orifices. Some pollen may possibly be thrown upon the single small stigma at the tip of the style, which rises much above the stamens. But the anthers are not dislodged when undisturbed, at least until after the elasticity of the filaments is lost : they are dislodged by humble-bees, which circle on the wing over the blossom, the FIG. 455 Vertical section of a flower-bud of Kalmia latifolia, showing the anthers lodged in the pockets of the corolla. 456. Expanded flower, with bowed stamens. 457. Vertical section of the same. 458. A stamen, enlarged. 230 THE FLOWER. under side of the abdomen frequently touching the stigma, while the proboscis is searching round the bottom of the flower, liberat- ing the stamens in the process, which one by one project their pollen upon the under side of the insect's body. In the passage from (lower to (lower, pollen is thus conveyed from the anthers of one to the stigma of another. 420. Iris has three stamens, one before each sepal or outer lobe of the perianth, and behind each petal-like lobe of the st\ le (Fig. 4-VJ) : the stigma, a shelf-like plate of each lobe, is just above the anther; but, as the anther faces outward and the stigma is higher and laces inward, no pollen can find its way from the one to the other. But the adaptation of parts is admir- able for conveyance by bees, which, standing upon the only landing place, the re- curved sepal, thrust the head down below the anther, and in raising it carry off pollen, to be afterwards lodged upon the stigmas of other flowers which they visit. 421. Transportation of Pollinia, or of all the pollen in a mass, is effected in most of the species of two large orders, not otherwise allied, the Asclepiadacca- and the Orchidacej-e. AVhile in the Iris family the number of stamens is reduced from six to three, in all the Orchis family, except Cypripedium, the stamens an- further reduced to a single one; but the pollen is peculiarly economized. That of Arethusa is in four loose and soft pellets, in an inverted casque-shaped case, hinged at the back, resting on a shelf, the lower (ace of which is glutinous stigma, over the front edge of which the casque-shaped anther slightly projects : and this anther is raised by the head of a bee when escaping out of the gorge of the flower. The loose pellets of pollen are caught upon the lice's head, to the rough sur- face of which they are liable to adhere lightly and so to be carried to the flower of another individual, there left upon its glutinous FIG. 459. Flower of Iris pumila. with front portion and half of one petalnid style- lobe and stij;iii:i cut away. The section of the stigma is seen edgewise: the rough upper surfare only is stigmatic. ADAPTATIONS FOR INTERCROSS I Mi. 231 stigma by the same upward movement which immediately after- ward raises the anther-lid and carries away its pollen, to be transferred to a third blossom, and so on. 422. But it is in Orchis and in the commoner re- presentatives of Orchis in North America (viz. Ila- benaria, &c.) that the most exquisite adapta- tions are found, and the greatest economy se- cured ; paralleled, how- ever, by most of the very numerous and various epi- phytic and by various ter- restrial Orchids of warmer regions. A single illus- tration ma^' here suffice ; and Darwin's volume on the Fertilization of Or- chids (396, note), with its references to the copious literature of the subject, may be studied for full particulars and their bearings. The flower is trimerous, and the peri- anth adnate to the ovary, therefore apparently de- veloped upon its sum- mit. The three external parts of the perianth, which in Ilabenaria orbi- culata (Fig. 460) are much the broader, are the sepals: the three alternate and internal, the petals : the base of the long and narrow petal which is turned downward is hollowed out and extended below into a long tube, closed at bottom, open at top (the spur or nectarj'), in which nectar is FIG. 460. Flower of Habenaria or Platanthera orbiculata, enlarged. 461. Combined stamen and stigma, more enlarged. 462. One of the two pollen-masses (pollinia). \vitb its stalk and glutinous disk or gland. 462". Lower part of this stalk and its disk, more magnified. THE FLOWER. copiously secreted and contained. The central part of the blossom, beyond the orifice of the nectary (shown separately in Fig. 461), consists of one anther and a stigma, fused together (the clinandrium) : the marginal portions, opening by a long chink, are the two cells of the anther, approximate at their broader portion above, widely divergent lielow : most of the lower part of the spaee between is cxces>ively glutinous, and is the stigma. The grains of pollen are united by means of short threads of very elastic tissue into small masses, and these into larger, and at length into pellets, having stalks of the same elastic tissue, by which they are all attached to a firmer central stalk, or cuudicle. (Fig. id.".- 1(!5.) To the lower end of this caudicle (directly to the end of it in our Ilabenariae and Orchises gener- ally, in this instance to the inner side of tin1 end. with a thick inter- mediate base intervening), is at- tached a button-shaped disk, the face of which is exposed, and is on a line with the surface of the anther ; so that these two disks look toward each other across the broad intervening stigmatic space, as seen in Fig. -li'il. The exposed face of the disk being covered with a durable layer of very viscid mat- ter, the body itself is sometimes termed agland, andnot impro] >crly . The viscidity is nearly of the same nature as that of the interven- ing stigma, of which the glands are generally supposed to be detaeheil portions. 1 1' so, then a portion of the stigma is cut off from the rest and specialized to the purpose of eonveyauee of the pollen. When a linger's end or any smaller body is touched to these disks, they adhere so firmly that the attached /x>ll!n!f/,i<»is dimorphism (4:5-1) the intent to self-fertili/e is evident. There may also be dimorphism as to the perianth, not partictilarlv affecting fertilization. One kind, however, and the commonest, is a special adaptation to intercrossing, viz. : li'l. Hetcrogonous Dimorphism. (11:5. note.) This term is applied to the case in which a species produces two kind> ol hermaphrodite flowers, occupying different individuals, the llowero essentially similar except in the andm-cinm and gynu-cium. l.iu Ihese reciprocally different in length or height, and the adapta lions such that, by the agency of insects, the pollen from the stamens of the one sort reciprocally fertilizes the stigma of tin- other.2 This dimorphism has been detected ill about fort v genera belonging to fourteen or fifteen natural orders, widely scattered through the vegetable kingdom ; but there are far more examples among the Rubiaceas than in any other order. Sometimes all the species of a genus are heterogonous, as in Honstonia. and The reported sensitiveness of the gland, referred to in the first issue of this volume (1S7!>), was founded upon misinterpreted observations. 1 This peculiar ;irrangoment has been long known in a few plants, such as Primula veris. 1'. grandiflora. and Houstonia. In Torrey and Grav's Flora of North America, ii. :',8. :V.t ( ls|:!). these (lowers are said to IK- diu-cio- dimorphous, not denoting that they an- at all unisexual, hut that the two forms occupy different individuals. Their meaning was detected by C. Darwin, and made known in his paper "On the Two Forms or Di rphiu Condition in the Species of Primula, and on their Kemarkable Sexual IMa- tions." published in the Journal of the Limiean Society, vi. (IS(L'), 77: repub- lislied, in 1S77. as tin- leading chapter of his volume entitled "The Different Forms of Flowers on Plants of the Same Species." Mr. Darwin had termed these flowers simply fti',,i,ir/,lii,- ; but in this volume he adopted Ililde- brand's name of //, t, ,;,*/„/,,/ for this kind of blossom. The difference, however, affects the audnivium, and even the pollen, as well as the style; when-fore we proposed for it the name of //ut if the organization wen- of three forms, any two of which inter- crossed with perfect fertility, the chances (as Darwin remarks) are two to one that any two plants were of dillereiit forms, and therefore by fertilizing each other completely fruitful. 428. The earliest known instance of three forms as to recip- rocal relative length of stamens and pistil is that of Lythrum 1 Impotence of own pollen, either absolute or relative, occurs no less in certain tlowers which are not dimorphous, as in Corydalis, some species of Passiflora, &<•. On the contrary, many dimorphous flowers are in a certain decree self-fertile, especially in the lonij-stamcncd and short-styled form. These subjects arc physiological, and belong to another volume. FIG. 468. Loiig-styliwl flower of Fig. 4f>7. laid open. 469. Long-stain ened flower of the panic laid open I'.olli equal] ADAPTATIONS FOE INTERCROSSING. 237 Long- styled. Mid- styled. Salicaria. This was indicated by Vaucher in 1841, more par- ticularly described by Wirtgen in 1848, but was interpreted by Darwin, and the more recondite differences brought to notice, in 1864. 1 "The three forms may be conveniently called, from the unequal length of their pistils, the long-styled, mid-styled, and short-styled. The stamens also are of unequal lengths, and these may be called the longest, mid-length, and shortest." The pollen of the different classes of stamens is of two sorts as to color, and of three as to size, the largest grains from the largest stamens. "The pistil in each form differs from that in either of the other forms, and in each there are two sets of stamens, different in appearance and func- tion. But one set of stamens in each form corresponds with a set in one of the other two forms. Altogether, this one species includes three females or female organs, and three sets of male organs, all as distinct from one another as if they belonged to different species ; and, if smaller functional differences are considered, there are five distinct sets of males. Two of the three hermaphrodites must coexist, and pollen must be carried by insects reciprocally from one to the other, in order that either of the two should be fully fertile ; but, unless all three forms coexist, two sets of stamens will be wasted, and the organization of the species as a whole will be incomplete. On the other hand, when all three hermaphrodites coexist, and pollen is carried from one to the other, the scheme is perfect : there is no waste of pollen and no 1 In an article On the Sexual Relations of the Three Forms of Lythrum Salicaria, in Jour. Linn. Soc. viii. 169. Also on the Character and Hybrid- like Nature of the Offspring of the Illegitimate Unions of Dimorphic and Trimorphic Plants. Ibid. x. 393, 1868. Reproduced and extended in his volume entitled "Forms of Flowers," 1877. FIG. 470. Diagram of the flowers of the three forms of Lythrum Salicaria, in their natural position, with the petals and calyx removed on the near side. The dotted lines with the arrows show the directions in which pollen must be carried to each stigma to ensure full fertility. (From Darwin.; Short- stvled. 470 238 THE FLOWKK. false coadaptation." The whole arrangement is displayed in the annexed diagram (Fig. 470 ). and in the following account of tlie operation.1 " In a state of nature, the flowers are inces- santly visited lor their nectar by hive and other bees, \arious Diptera. and Lepiiloplera. The nectar is secreted all round the base of the ovarimn : but a passage is formed along the upper and inner side of the Ilower by the lateral deflection (not repre- sented in the diagram) of the basal portions of the filaments; so that insects invariably alight on the projecting stamens and pistil and insert their proboscides along the upper and inner margin of the corolla. We can now see why the ends of the stamens with their anthers and the end of the pistil with the stigma are a little upturned, so that they may be bru>hed by tlie lower hairy surfaces of the insects' bodies. The shorie-t stamens, which lie1 enclosed within the calyx of the long- and mid-styled forms can be touched only by the proboscis and narrow chin of a bee: hence they have their ends more upturned. :md they are graduated in length, so as to fall into a narrow file, sure to be raked by the thin intruding proboscis. The anthers of the longer stamens stand laterally farther apart and are more nearly on the same level, for they have to brush against the whole breadth of the insect's body. . . Now 1 have found no exception to the rule that, when the stamens and pistil are bent, they bend to that side of the flower which secretes nectar. . . . When nectar is secreted on all sides, they bend to that side where the structure of the Mower allows the easiest access to it, as in Ly thrum. ... In each of the three forms, two sets of sta- mens correspond in length with the pistil in the other two forms. When bees suck the Mowers, the anthers of the longest stamens, bearing the green pollen, are rubbed against (lie abdomen and the inner sides of the hind legs, as is likewise the stigma of the long-styled form. The anthers of the mid-length stamens and the stigma of the mid-styled form are rubbed against the under side of the thorax and between the front pair of legs. And, lastly, the anthers of the shortest stamens and the stigma of the short -styled form are rubbed against the proboscis and chin; for the bees in sucking the flowers insert only the front part of their heads into the Ilower. On catching bees, I observed much green pollen on the inner sides of the hind legs and on the abdomen, and much yellow pollen on the under side of the thorax. There was also pollen on the chin, and, it may be presumed, on the proboscis, but this was difficult to observe. I had, however, 1 All from Darwin, Forms of Flowers, 137-147, &c. ADAPTATIONS FOR INTERCROSSING. 239 independent proof that pollen is carried on the proboscis ; for a small branch of a protected short-styled plant (which produced spontaneously only two capsules) was accidentally left during several da}7s pressing against the net, and bees were seen insert- ing their proboscides through the meshes, and in consequence numerous capsules were formed on this one small branch. . . . It must not, however, be supposed that the bees do not get more or less dusted all over with the several kinds of pollen ; for this could be seen to occur with the green pollen from the longest stamens. . . . Hence insects, and chiefly bees, act both as general carriers of pollen, and as special carriers of the right sort." 429. Finally, a long series of experiments (requiring eighteen distinct kinds of union) proved that both kinds of pollen are nearly or quite impotent upon the stigma of the same flower, and that no ovary is fully fertilizable by other than a u legitimate union," i. e. by stamens of the corresponding length ; but that the mid-length pistil is more prolific than either of the others under illegitimate union of either kind ; which might perhaps be expected, as the pollen proper to it is intermediate in size of grains between that of the long and that of the shortest stamens. 430. Nesaea verticillata, a common Lythraceous plant of the Atlantic United States, is similarly trimorphous, but has not yet been particularly investigated. Several South African and American species of Oxalis are equally trimorphous, and have been investigated by Darwin and Hildebrand,1 with results quite as decisive as in Lythrnm Salicaria. One genus of Monocotyledons has trimorphous blossoms, viz. Pontederia, of which the North American P. cordata is a good illustration.2 431. All known flowers exhibiting reciprocal dimorphism or trimorphism are entornophilous : no such wind-fertilized species is known. Few of them are irregular, and none very irregular : they do not occur, for instance, in Leguminosae, Labial a1, 1 Monatsber. Akad. Berlin, 1866; Bot. Zeit. 1871, &c. According to Darwin, Fritz Mueller " has seen in Brazil a large field, many acres in extent, covered with the red blossoms of one form [of an Oxalis] alone, and these did not produce a single seed. His own land is covered with the short-styled form of another species, and this is equally sterile ; but, when the three forms were planted near together in his garden, they seeded freely." Forms of Flowers, 180. 2 Detected by W. H. Leggett. See Bulletin of Torrey Bot. Club, vi. 62, 170; and for the original discovery in Brazilian species, by Fritz Mueller, see Darwin's Forms of Flowers, 183, &c. Pontederia has three lengths of style and counterpart stamens, as in Lythrum Salicaria, each flower having two sets of stamens, three in each set. 240 THE FLOWER. Scrophulariaceae, Orchidaceae, &c. Nature is not prodigal, and dot's not endow with needless adaptations flowers which are otherwise provided for. § 3. An AIM A i IONS FOR CLOSE FERTILIZATION. l.'ii'. Kveu when- cross-fertilization in bisexual flowers is obviously arranged lor. it is apt to be tempered with more or less of close-fertilization. The more exquisite the arrangements for the former are, the more completely is the plant dependent upon insect visitation. Failure to intercross is a remote and small evil compared with failure to set seed at all. In order therefore that the plan of cross-fertilization may not defeat even its own end, through too absolute dependence on precarious assistance, some opportunity for self-fertilization will usually be advantageous. Also there is a long array of insect-visited flowers, especially polyandrous ones, in which close fertilization must lie much the commoner result, except where the pollen of another but wholly similar flower has greater potency. 433. Subsidiary self-fertilization is secured in a great variety of ways. In Gentiana Andrewsii. which is proterandrons, and usually cross-fertilized by humble-bees entering bodily into the corolla, an exposed surface of pollen long remains fresh upon the ring of anthers girding the base of the style : when the stigmas separate, the}" remain for some da}*s simply divergent, but they at length become so revolute that the receptive surface is brought into contact with the ring of pollen below. The opening and closing of blossoms by day or night, the growth of style, fila- ments, or corolla after anthesis commences, or other changes of position, may secure a certain amount of self-fertilization in a subsidiary or even in a regular way. Then certain species, such as Chickweed. which blossom through a long season, close- I'ertilize even in the bud in early spring, when insects are scarce, but are habitually intercrossed by insects in summer. Somewhat similarly, according to Hermann Mueller.1 certain species, such as Euphrasia ..Micinalis and Khinanthus Crista-galli. habitually produce two kinds of blossoms, one larger and more showy, usually ailed ing sunny localities, and with parts adapted to intercrossing by insects : the other smaller or inconspicuous, and with anthers adjusted for giving pollen to the adjacent stigma without aid. There are gradations between these last arrange- ments, and the more special and remarkable one of dimorphism with — - 1 Befruchtung tier Blumen durch Insektcn, 294 ; Nature, viii. 433. ADAPTATIONS FOR CLOSE FERTILIZATION. "241 434. Cleistogamy. Here the intention and the accomplishment of self-fertilization are unmistakable. This peculiar dimorphism consists in the production of very small or inconspicuous and closed flowers, necessarily self-fertilized and fully fertile, in addition to ordinary, conspicuous, and much less fertile, though perfect flowers. Two cases were known to Linnaeus,1 and one of them to Dillenius before him ; those of Viola have long been familiar in the acaulescent species ; Adrien Jussieu made out the structure of the cleistogamous flowers in certain Malpighiaceae in 1832, and recorded in 1843 that Adolphe Brongniart had well investigated those of Specularia, and that Weddel had discov- ered them in Impatiens Nolitangere. A full account of the then known cases was given by Mohl '2 in 1863 ; but D. Mueller, of Upsala, who examined Viola canina, is said by Darwin to have given,3 in 1857, "the first full and satisfactory account of any cleistogamic flower." The appropriate name of cleistogamous was given by Kuhn,4 in 1867, and is now in common use. 435. Cleistogamous flowers are now known in about 60 genera, of between twenty and thirty natural orders, of very various relationship, though all but five are Dicotyledons. All but the Grasses5 and Juncus are entomophilous as to the ordinary flowers, and most of these such as have special arrangements for their intercrossing, either by dichogamy, heterogone dimorphism or trimorphism (in Oxalis) , or such special contrivances as those of Orchids. 436. It has been said that the ordinary flowers in such plants are sterile, and perhaps they always are so except when cross- fertilized : in most cases they are habitually infertile or spar- ingly fertile. Probably they suffice to secure in every few generations such benefit as a cross may give, while the principal 1 Campanula (now Specularia) perfoliata and Ruellia clandestina, the latter a cleistogamous state of R. tuberosa. Linnaeus did not make out the structure of the flowers, but supposed them to want the stamens. 2 In Bot. Zeitung, xxi. 309. 3 In Bot. Zeitung, xvi. 730. * Ibid. xxv. 65. The name (denoting "closed up "union or fertilization) has been written deistogenous, which is not so proper. We prefer dei'slot/amou* to cleistogamic (and so of similar terms), as best harmonizing with tin- Latin adjective form, both in form of termination and in euphoniously taking tin- accent upon the antepenult. 5 Amphicarpum (Milium amphicarpon, Pursh) is the earliest recognized cleistogamous Grass, except perhaps Leersia oryzoides. Some species of Sporobolus are like the latter, and Mr. C. G. Pringle has recently detected such flowers concealed at the base of the sheaths in Danthonia. Amer. Jour. Sci. January, 1878, 71. 16 •J4- THE FLOWER. increase is by clcistogamous self-fertilization, which thus offsets the incidental disadvantage of the former mode. 437. In general, the cleistogamous arc like unto the ordinary flowers arrested in development, some arrested in the almost fully formed lmd, most at an earlier stage, and in the best marked cases with considerable adaptive modification. In these, "their petals are rudimentary or quite aborted; their stamens are often reduced in number, with anthers of very small size, containing few pollen-grains, which have remarkably thin transparent coats, and generally emit their tubes while still enclosed within the anther-cells ; and, lastly, the pistil is much reduced in size, with the stigma in some cases hardly at all developed. These flowers do not secrete nectar or emit any odor : from their small size, as well as from the corolla beiii" o rudimentary, they are singularly inconspicuous. Consequently, insects do not visit them ; nor, if they did, could the\* find an entrance. Such flowers are therefore invariably self- fertilized ; 3ret they produce an abundance of seed. In several cases, the young capsules bury themselves beneath the ground, and the seeds are there matured. These flowers are developed before, or after, or simultaneously with the perfect ones." 1 In Grasses, however, as in some Dicotyledons, there is much less modifica- tion and more transition. For when Leersia half protrudes its panicle, in the usual way, the included half is fertile and the expanded portion sterile (or almost alwa3Ts so), although the flowers may open and exhibit well-developed anthers, ovaries, and stigmas. But when similar panicles remain enclosed in the leaf-sheaths, they are mostly fruitful throughout. 438. Fully to apprehend the economy of cleistogamy in pollen- saving alone, — and contrariwise to estimate the expense of intercrossing. — one should compare the small number of pollen- grains which so completely serve the purpose in a typical cleis- togamous llower (say Ion in ( )\alis Acetosella. ^."iii in Impatiens, 100 in some Violets) with the several thousands of all entomo- philons cross-fertilized (lowers, rising to over three and a half millions in the llower of a Peony, also their still greater number in many aneniophilous blossoms. To this loss should be added (lie cost of ;i corolla and its action, also of the production of odorous material and of nectar. No species is altogether cleis- togamous. Thus cleistogamy, with all its special advantage, testifies to the value of intercrossing. 1 Darwin, Forms of Flowers, 310. PERIANTH, OR FLOWER-LEAVES. 243 SECTION V. THE PERIANTH,* OR CALYX AND COROLLA IN PARTICULAR. 439. The distribution of the floral leaves around the axis, which belongs to phyllotaxy, and their particular disposition in the bud (aestivation), have already been considered in Chap. IV. Sect. I., II. And most of the morpholog}' of cabyx and corolla has been outlined in the preceding sections of the present chap- ter. What remains chiefly relates to particulars of form and to terminology. 440. Duration. The differences in this respect give rise to a few terms, such as the following. Calyx or corolla may be Persistent, not cast off after anthesis, but remaining unwithered until the fruit is formed or matured ; as the calyx in Labiatse, in Physalis, and most Roses. Marcescent, withering or drying without falling away ; as the corolla of Heaths, Drosera, &c. Deciduous, falling after anthesis and before fructification ; as the petals of Roses, the calyx and corolla of Columbine. Ephemeral or Fugacious, lasting for only a day ; as the petals of Poppy, Helianthernum, Purslane, and Spiderwort. In the two former, they are cast or early deciduous, the anthesis lasting but a day : in the two latter, the anthesis is equalby or more brief, but the petals deliquesce or decay at once without falling, as does the whole flower of Cereus grandiflorus and other night- blooming Cactaceas. Caducous, falling when the blossom opens ; as the calyx of Poppy and Baneberry. 441. Numerical Terms, succinctly denoting the number of leaves, either of the perianth as a whole, or of any one of its circles, are common in descriptive botany. The most general are those which simply specify the number of component leaves, by prefixing Greek numerals to the Greek name of leaves, ex- pressing them in Latin form, or transferring them to the Eng- lish. Thus Diphyllous, of two leaves (sepals or petals) ; Triphyllous, of three ; Tetrfip/>yllous,offour ; Pentaphylloits. of five ; Hcxaphyllous, of six, and so on. A tulip and a Tradescantia flower have a hexaplryllous perianth, but composed of two circles, answering to calyx and corolla ; each TriphyUous.z When the character 1 Perianthium, alias Per/i/one or Perigonium. (296.) 2 As elsewhere explained, when numerical composition is indicated without reference to nature of parts, the terms dimerous, trirncrous, tetramerous, penta- merous, &c., may be used. (322.) 244 THE FLOWER. of the organ, i. e. whether calyx or corolla is to be specified, the word sepal or petal is employed in the combination ; as. Disepalous, of 1 \\ <> .sepals ; Trisepalous, of three ; Tetrasepulous, of four ; Pentasepalous, of five (also written 5-sepalous, and ac- cordingly 2-sepalous, 3-sepalous), and so on: also, Dlfx'inlnus, Tripetalous, Tetrapetalous, Pentapetalous (2-5- petalous), &c., when the corolla is concerned. 412. Monophyttous, Monosepalous, and Monopetalous are the proper terms for perianth (calyx, corolla, &c.) composed of a single leaf. Likewise PolyphyUous, Potysepalous, and JJ<>/>/j>r/i//- ous for the case of a considerable but unspecified number of members. Unfortunately, in the Linua-an and long-prevalent use, monopetalous was the term employed to designate a corolla of one piece iii the sense, or the fact, of a coalescence or grow- ing together of two, three, five, or more petals into a cup or lube ; and so of a calyx, of a whorl of bracts, &c. And poly- petalons, polysepalous, and polyphyllous were the counterparts of this, meaning of more Mian one distinct piece, whatever the number. The misleading use, consecrated b}' long prescription, is not yet abandoned, but will in time be obsolete. In present descriptive botany, a polyphyllous calyx, or a polypctalous corolla, or a 5-petalons corolla, would be taken to mean that the sepals or petals (as the case may be) were distinct or uncom- bined, and a monopetalous corolla to be one with petals combined by coalescence. (.'J2'.).) 443. Terms of Union or Separation. The proper term for a corolla or a calyx the leaves of which are more or less coalescent into a cup or tube is Gamopetalous for such a corolla, Gamosepalnns for the calyx : these terms meaning united petals or sepals. The older and mis- leading names Mbnopetalous &nd Monosepalous, although current up to a recent day, should be discontinued. Another term is not rarely used in (lermany, that of Si/ni/x-fn/mts. for the gamo- petalons (or formerly nionopetalons) corolla. --therefore .s////- SI'/HI/IIHS for a similar calyx. It is perhaps a more apt term than gamopeta lous, and of the same etymological signification; but the latter is already well in use. Choripetalous is, on the whole, the most fitting name for a corolla the petals of which are separate' (as it literally expresses this), that is, for what is still commonly called Polypetalous, as already explained. ( M2.) It is adopted by Eichler, &c. C/tori- eepalous is the term applied to the calyx. Diali/petalous (em- ployed by Kndlicher) has the same meaning. Both this term and choripetalous carry the implication of separated, rather PERIANTH, OR FLOWER-LEAVES. 245 than of typically separate, parts. Eleutheropetalous (literally free-petalled) has also been used, but is inconveniently long. 444. Degree of coalescence is most correctly expressed by the phrases united (connate, or coherent, or coalescent) at the base, to the middle, or to the summit, as the case may be. But it is more usually and tersely expressed in botanical description by employing terms of division, identical with those used in describ- ing the lobing or toothing of leaves and all plane organs. (184-188.) That is, the calyx or corolla when gamophyllous is for description taken as a whole, and is said to be parted (^-parted, 5-parted, &c.), when the sinuses extend almost to the base ; cleft, when about to the middle ; lobed, a general term for any considerable separation be}rond toothing ; dentate or toothed (%-toothed, ^-toothed, &c.), when the union extends almost to the summit ; entire, when the union is complete to the summit or border. 445. Parts of Petals, &c. The expanded portion of a petal, like that of a leaf, is the LAMINA or BLADE : any much contracted base is the UNGUIS or CLAW. The latter is very short in a rose- 474 petal, but long and conspicuous in a pink and all flowers of that tribe (Fig. 471), in many Capparidese (Fig. 409) and Cruciferse. A sepal is very rarely distinguishable into lamina and claw. 446. Parts of Gamophyllous Perianth. The coalescent portion of a corolla, calyx, or of a perianth composed of both (such as a Lily or Crocus-blossom) , so far as the sides are parallel or not too spreading, is its TUBE : an expanded terminal portion, either divided or undivided, is the LIMB or BORDER. The limb may FIG. 471. Corolla of Soapwort. of five separate long-clawed or unr/uiculate petals, with a crown at the junction of claw and blade. FIG. 472. Flower of Gilia coronopifolia ; the parts answering to the claws of the petals of the last, figure here all united into a tube. FIG. 473. Flowerof the Cypress- Vine (IpomceaQuamoclit); the petals a little farther united into a five-lobed spreading border. FIG. 474. Flower of the Ipomrea coccinea; the five component petals perfectly united into a trumpet-shaped tube, and beyoud into an almost entire spreading border. 246 THE FLOWER. be parted (that is, the component parts not united) quite or nearly down l<> tin- tube or base, as in Fig. 171'. 17."> ; or less so, as in Fig. 17.".. 17G (with limb o-lobed) ; or with merely angles or points to represent the tips of the component members, as in Fig. 471 : or with even and entire border, as in common Morning- Glory, Fig. 4*-2. 447. The line, or sometimes a manifest or conspicuous portion, between the limb and tube (in the corolla always a portion above or at the insertion of the stamens, when these are borne by the corolla) is called the TIIKOAT, in Latin FAUX, pi. fauces. This is mostly more open than the tube, yet less expanded than the limb : but it often presents insensible gradations from the- one to the other. 448. Such appendages as the CORONA or ( KOWN (:'..sf>, shown in Fig. K»:i, 404, 471) usually belong to the throat of a gamo- petalous corolla or perianth, as in Oleander, Comfrey, Borrage, Narcissus, &c., or to a corresponding position when the parts are not coalescent. 449. Forms of Corolla, Calyx, &c. As to terminology, some of these are special and are applicable to corolla only, as the Papilionaceous , the peculiar irregular corolla of the typical portion of Leguminosae (388, Fig. 342-334). which has been already illustrated, and in which the petals, two pairs and an odd one. take particular names. Also the Caryopkyllaceous, or Pink-flower (Fig. 171 ). a regular corolla, of five long-clawed ( tt/n/i/ir/i/nff ) petals, the claws enclosed in a tubular calyx and the blades spreading; and the Cruciferous, of (bin somewhat similar petals, the four abruptly spreading blades in the form of a cross (c,-u<'/'nte), as in Fig. 394. Rosaceous, with roundish and widely spreading petals on very short or hardly any claws, as in Rose and Apple-blossoms. /./////><)- craterium, which the name refers to, with a stem or handle beneath, is now to be met with only in old pictures. 7'n/i/i/fir. when strictly used, denotes a gamophyllous perianth with limb inconspicuous in proportion to the tube, as in Trumpet Honeysuckle, or as Fig. 472-471 would be if the limb were much diminished or wanting. But it is some- times used in the sense of having a conspicuous tube. Kl')!. Tetniili/iitnnoiis is similarly applied to that of six stamens, two of them shorter, in the manner characteristic of Cruciferae, Fig. 396. !.">!. Terms which denote coalescence of stamens, whether by their filaments or their anthers, are Mnniiilfljthous^ that is, in one brotherhood, by coalescence of the filaments into a tube, as in the Mallow (Fig. 48.")). Lupine (Fig. 484), Lobelia (Fig. 488), &c. D!titl<-li>li»u.s. in two brotherhoods, by coalescence of the fila- ments into two sets; sometimes an equal number in each, as in Fumariaceae (Fig. 390), sometimes nine in one set and one separate, as in the Pea (Fig. 483) and most Papilionacea-. Triadelphous, with filaments united in three sets or clusters, as in Ilypericum. Pentadelphous, in five sets, as in Linden, Fig. 398, 399. But in general, when the sets are several, without regard to the number the stamens are said to be Polyadelphous. Syngenesious, when the stamens are united by their anthers into a tube or ring ; as in the whole vast order of Composite •• stig. •is: i 191 m ( Fig. 486, where they are five in number and the filaments dis- linct), in Cuciii-bita (Fig. 489, 490, where they arc three in number and the filaments partly monadelphous), and in Lobelia (Fig. INN, where the}- are also five and the long filaments are mainly monadelphons) . Kid. Iss. Flower c,|" Lobelia eanlinalis. with tnhr of cor., lla divided mi our side; filaments and anthers united into a tube: /! tube of filaments; ".of antin-rs. I-'li;. !-!> Male llower of Cm-ui-liita (Squash), with liinli of calx \ ami rorolla cut away, to show Ihr stamens, \\7... tlir.'c tilainonts. scparatr at luisc luit monadelphOUS al'ovc, ami tlitvr synu'cncsioiis anthrrs in a kind of licail. 4!H». Staim-lis of the same, fiilart;cil ami tlir HI>IM'V I'art cut away, to show the union. Tin- anthers are sinuous. 491. A ili'tadn-il stamen of the Melon, with loosely sinuous anther. I-' 11,'. l:ij. Stamens ami style of a C'yiirii>ciliuin. united into one body or column: O. anthers; »7. enlarged sterile stamen ; .s/ir/. the stigma. ANDKCEC1UM, OK STAMENS. 251 455. Of terms relating to adnation of stamens, besides the general ones of hypogynous, perigynous, epigynous (332), and eplpetalous, or adnate with corolla, there is the special one of Gynandrous, having stamens borne upon the pistil, as in Orchidacese. In Cypripediurn, the filaments of two stamens, and an enlarged sterile stamen behind, are adnate to a style, while the two anthers are quite free (Fig. 492) ; in the proper Orchis tribe (as in Fig. 460, 461), anther and stigma are consolidate] into one mass, and there is no evident style. 456. A complete stamen consists of FILAMENT and ANTHER. The latter is the functionally essential part of the organ, and therefore is wanting only in abortive or sterile stamens. (345, 352, &c.) The filament, being only a stalk or support, may be very short or wholly wanting : then the anther is sessile, just as the blade of a leaf is said to be sessile when there is no petiole. 457. The Filament, although usually slender and stalk-like, assumes a great variety of forms : it is sometimes dilated so as to resemble a petal, except b}' its bearing an anther ; as in the transition states between the true petals and stamens of Nyrn- phsea, shown in Fig. 318. 458. Such petaloid filaments would indicate that this part of the stamen answered to blade rather than to footstalk, while others would harmonize better with what seems at first sight to be the more natural view, that the filament is the homologue of the petiole, the anther of the blade of a leaf. Remembering that in large numbers of leaves there is no distinction into petiole and lamina or blade, such homologies should not be insisted on. The filament may be variously appendaged by outgrowths. Some of these appendages are very conspicuous, such as the scale of Larrea (Fig. 405), which is on the inside, and the nectariferous hood of Asclepias on the outside ; or there may be a tooth on each margin, as in species of Allium. 459. The Anther, the essential organ of the stamen, contain- ing the pollen, surmounts the filament, when that is present. It normally consists of two cells or lobes, the word cell being here used in the sense of sac. But, as each sac is not rarely divided into two cavities (locelli), the best technical name for anther-sac is that of THECA. The two thecse, lobes, or cells are commonly connected by a more or less evident and sometimes conspicuous common base or junction, which is mostly a pro- longation of the filament, the CONNECTFVUM, or in English CONNECTIVE. 460. For the discharge of the pollen, the cells of a normal anther open at the proper time by a line or chink, usually 252 THE FLOWER. extending from top to bottom (Fig. 493), the suture or line of dehiscence. Commonly this line is lateral or marginal : not rarely it laces forward or backward. In the vast genus Solanum, 493 to which the Potato belongs, in most Ericaceous plants (Fig. 458, 494), in Polygala, and in many other flowers, the anther- cells open only by a hole (foramen or pore} , or at most a short chink, at the tip, through which the pollen has in some way to be discharged. In Vaccinium (Cran- berry, Bluebeny, &c.), the pore-bearing tip of the anther-cell is prolonged considerably, often into a slender tube, as in Fig. 340. In the Barberry (Fig. 495) and in most of that family, also in Lauracets, 499 the whole face of each anther-cell separates by a con- tinuous line, forming a kind of door, which is attached at the top, and turns back, as if on a hinge : in this case, the anthers are said to open by uplifted valves. In the Sassafras and many other plants of the Laurel family, each lobe of the anther opens by two smaller valves of the kind, like trap-doors. 4G1. The attachment of the anther to the filament presents Hi ice principal modes, which are connected by gradations. These are the Innate (Fig. 495, 49G), in which the anther directly continues and corresponds to the apex of the filament, the cells usually dehiscent strictly marginally, the lobes or cells not looking or projecting either inward or outward. 1<'I<; in:;. \ stamen, \viii, its anther. /<, surmounting the filament, a, and opening in (ho normal manner ilown the whole length of the outer side of each cell. FIO. 494. Stamen of a Pyrola; each cell of the anther opening by a terminal ori- fice or pore. FIG. 495. Stamen of a Barberry; the cells of the anther opening each by an up- lifted valve. FIO 406. A stamen of Isopyrum biternatum. with innate anther. 407. Stamen of Liri-idendron, or Tnlip-trcc. with adn.ito extrorse anther. 498. Stamen of CEnothera Rlauea, with the anther fixed l>y its middle and versatile. FIO. 499. A stamen of Asarum Canadense, with adnate anther and prolonged tip to connective. ANDRCECIUM, OR STAMENS. 253 Adnate, in which the connective appears to be a direct con- tinuation of the filament, having the anther adherent to the anterior or posterior face of it, and the lines of dehiscence therefore looking inward or outward. Magnolia, Liriodendron (Fig. 497), and Asarum (Fig. 499) furnish good examples ; the latter conspicuously so, on account of a prominent prolongation of the connective or tip of the filament. Versatile, when the anther is attached at some part only of its, back or front to the tip of the filament, on which in authesis it lightly swings ; as in Plantain, in all Grasses, the Lily, Evening Primrose (Fig. 498), &c. 462. The direction to which an anther faces, whether inward (toward the centre of the flower), or outward (toward the peri- anth) , has to be considered ; except in the case of an innate anther with strictly lateral or marginal dehiscence. An anther is Extrorse, i. e. turned outward, or Posticous, when it faces to- ward the perianth, as in Magnolia and Liriodendron (Fig. 497), Asarum (Fig. 499), and Iris ; these all being cases of adnate and extrorse anthers, the cells attached for their whole length to the outside of the summit of the filament or the connective. Introrse, i. e. turned inward, or Anticous, when it faces toward the axis of the flower ; as in Nymphaeacese (Fig. 318), in Violet and Lobelia (which are adnate and introrse), and in CEnothera. In the common Evening Primroses (as in Fig. 498) the anther is fixed near the middle, introrse, and versatile. 463. The direction in which the anther may be said to face, outward or inward, depends upon two characters, which do not alwa}-s coincide, viz. the insertion or attachment of the cells, and the position of their line of dehiscence. In such a strongly characterized adnate anther as that of Liriodendron (Fig. 497), both the attachment and the dehiscence are plainly posticous or extrorse : in most species of Trillium, the cells are introrse as to attachment, but some are nearly marginal and some are even rather extrorse as to dehiscence : in the related Medeola, and in Lilium, where the anthers are extrorsely affixed toward the base or middle to a slender tip of the filament, the dehiscence is either introrse or nearly marginal. Parnassia is in similar case ; the anthers being clearly extrorse as to insertion and more or less introrse as to dehiscence. 464. Adnate anthers are perhaps as frequently extrorse as introrse. Others, whether basifixed or medi fixed, are more com- monly introrse. Those fixed by the middle, or at any other part of the back, and lying on the inner side of the filament, are said to be Incumbent. 254 THE FLOWER. 465. The connective may be appendaged either by a prolon- gation or otherwise from the tip (as in Fig. -llt'J), or from the back, as in Violets and in many Kricaceoiis plants. I (ill. The normal anther is two-celled, bilocular, or (to use a less eoiiinion term) f/ifhecous, and its lohes or cells parallel, right and left : but the cells at first, and sometimes at maturity, are l>!l each is divided into two by a partition which stretches from the connective to the suture or line of dehiscence. In an innate anther, and in main' others, this line of dehiscence is marginal or lateral, either strictly or nearly so, as in Fig. 500. When introrse or extrorse (as in Fig. 5()1, 502), the sutures may still be considered to represent the margins turned inward or outward. The pollen is accordingly pro- duced in four cavities or separate portions of the interior. But *he two locelli on the same side of the midrib or connective (right and left) are usually confluent into one pollen-tilled cavity or cell at maturity if not earlier, or at least the partition between • hem breaks up at dehiscence. Sometimes it remains, and, the groove at the sutures being deep, the anther is strongly four- lobed or quadrilocular at maturity, as in Menispermum (Fig. 504) ; but morphologically this is still only bilocular (dithecous) although <|uadrilocellate, and the anther opens at the sutures and through these partitions. 467. A stamen being the homologue of a leaf, the natural supposition is that the anther is homologous with the blade or an apical portion of the blade, therefore the two lobes or thcca- with the right and left halves of it. the intervening connective with the midrib, and the line of dehiscence with the leaf- mar- gins.1 This conception is exemplified by the accompanying 1 'Phis is the view lonir airo taken by Cassini and Ku'per. and it may still lie maintained a- the best morphological conception. Mohl interposed some dltjertiniis in its universality: luit, as presented in Sadie's Text-Hook, they are not incompatible with tin1 common morphology. Sachs takes the fi la- ment with the connective to be the liomoloL'ue of the whole leaf, and the anther-cells as appendages. Others, in likening the anthers to glands, adopt a similar vie \v. FIC. .".on. I mint i- aiitlicr. smiie us Kit,'. l!H-.. in \ ounu'rr -t:iti-. \\illi transverse section, sliowiii'j tlir f'Hir li'ivlli. r.ni. Same of an adn.'ite extrorso antlir-r. snc-li as Fig. 407. ">()?. S:inir :is the priM-ciliiiu' luit mature ainl ilehisccut, the two locelli becoming one cell l>y the vanishing <>r lireaking up of the partition. ANDECECIUM, OR STAMENS. 255 diagram, Fig. 503, which should, however, show the median partitions in the cross-section, or traces of them. Pollen is a special development into peculiar cells of what would be parenchyma in a leaf. Its formation normally begins in four places, which may remain separate up to maturity, or the two on each side of the axis or connective may early be confluent into one cell. 468. Of the man}' deviations of the typical two- celled anther, with its cells parallel and united longitudinally by a connective, the simplest and commonest is that in which (as in Fig. 505) the two cells diverge below and remain united only at their apex. Next, the two cells ma}-, in their early development, become confluent at the apex, as in the Mallow family (Fig. 50G), so as to form a continuous pollin- iferous cavity within, opening by a continuous suture round the margin : here the anther is unilocular or one-celled b}T confluence- In another wa}', the anthers of some species of Orthocarpus (gew erally resembling Fig. 505, but the lobes or cells quite distinct or even separated at apex) lose one of the cells by partial c»* complete non-development and so become one-celled by abor tion. The anther of Gomphrena (Fig. 507) is completely un> locular b}T abortion or suppression of the companion cell. Thuu losing one half, it is said to be dimidiate, or halved. 469. The two anther-cells, such as those of Fig. 505, some- times diverge so much that they form a straight line transverse to the filament, as in Monarda (Fig. 508), in which their contiguous ends so coalesce as to give the appearance of a one-celled anther fixed by the middle. Or, again, the two cells may be separated by the enlargement of the connective between them, as in Cal- amintha, Fig. 509. This enlargement is extreme in the great genus Salvia. in which a very long and narrow connec- tive gives the appearance of a filament astride the apex of the FIG. 503. Diagram to illustrate the morphology of the stamen, on the idea that the anther answers to leaf-blade: the lower portion being filament and a part of the anther, in section, the upper a part of a leaf. FIG. 504. Stamen of Menispermum Canadense, the quadrilooellate anther divided. FIG. 505. Stamen of Pentstemon ptibescens, with anther-cells divergent. FIG. 506. Stamen of Mallow (one of the cluster of Fig. 485). the two cells and sutures confluent into one. FIG. 507. Anther ef Gomphreua or Globe Amaranth, meditixed, of a single cell, dehiscent. 256 Till; FLOWER. proper filament, and bearing an anther-cell at each end. In a few species, the two anther-cells arc nearly alike ; in more, the lower one is imperfect, as in Fig. 510°; in more, it is abortive or wanting altogether, as in Fig. 510fc. Then, in the related California!) genus Aiuliliertia. the lower half of this connective is reduced to a short tail, as shown in Fig. 511°, or even in most of the species to so minute a vestige that, except for these transitions, the stamen might be supposed to consist of a simple filament, with an interruption like a splice in tin- middle, and surmounted by a one-celled anther, as shown in Fig. all6. In Rosemary, the continuity is complete, although a minute reflexed tooth sometimes indicates the junction. 470. Polleii, the product of the anther, is usually a powdery substance, which when magnified is seen to consist of separate grains, of definite size and shape, uniform in the same plant, .-,11' 513 M4 I' but often very different in different species or families. The grains are commonly single cells, globular or oval in shape, and of a yellow color. But in Spiderwort they arc oblong; in the KKS. nos -r.ll. Anthers, with nppor pnrt of filament. «>f several Lahiata'. .r>OS. Of Monarda. ->n:i of. -i <':il:iiniiiihii. 510 Of two species of SaMa, ^wlth long and slender connective, the upper fork c,f which bears one anther-cell; the lower in a (from Snlvia Texanai. l.eaiiie,' the othercell in an imperfect condition; in l> (from S. coccinea), bear- ing none at all. nil.rc. Same of Audil.ertia irranditlora. the lower fork of the connec- tive reduced to a naked spur; l>. from A. stachyoides, in which this lower fork is nearly wanting, and the upper is in a straight line with the filament which it seems to continue. Kit;. 512-515. Forms of pollen: 512, from Mimulus moschatus; 513, Sicyos; 514, Kchinocystis; 515, Hibiscus. POLLEN. 257 Cichory and Thistle tribes, many-sided ; in the Musk-plant, spirally grooved ; in the Mallow family and the Squash and 517 521 Pumpkin, beset with briskly projections, &c. The pollen of Pine, as well as that of the Onagracese, is not so simple, but appears to consist of three or four blended cells ; that of most Ericaceae evidently consists of four grains or cells united. (Fig. 512-521.) The most extraordinary shape is that of Zostera, or the Eel-grass of salt-water, in which the grains (destitute of the outer coat) consist of long and slender threads, which, as they lie side by side in the anther, resemble a skein of silk. 471. Pollen-grains are usually formed in fours, by the division of the living contents of mother cells fu\st into two, and these again into two parts, which become specialized cells. As the pollen completes its growth, the walls of the mother cells are usually obliterated. But sometimes these cells persist, either as shreds, forming the cobweb-like threads mixed with the pollen of Evening Primrose, or as a kind of tissue combining the pollen into coherent masses, of various consistence. Of this kind are the elastically coherent pollen-masses (or POLLINIA, sing. POLLINIUM) of Orchises (Fig. 463), and the denser waxy ones of mairy other orchids and those of Asclepias or Milkweed, Fig. 522. 472. A pollen-grain has two coats. The outer coat is com- paratively thick, and often granular or fleshy. This is later formed than the inner, and by a kind of secretion from it : to it all the markings belong. The inner coat, which is the proper cell-wall, is a very thin, delicate, transparent and colorless mem- brane, of considerable strength for its thickness. The pollen of Zostera and of some other aquatic plants is destitute of the outer coat. 473. The cavity enclosed by the coats is filled with a viscid substance, which often appears slightly turbid under the higher powers of ordinary microscopes, and, when submitted to a mag- FIG. 516-521. Forms of pollen: 516, Lily; 517, Cichory; 518, Pine; 519, Circrea; 520, Kalmia; 521. Evening Primrose. FIG. 522. A pair of pollinia of Asclepias, annexed by their caudicles to the gland. 17 258 THE FLOWER. inlying power of about three hundred diameters, is found to contain a multitude of minute particles (fovillce) , the larger of which are from one four-thousandth to one five-thousandth of an inch in length, and the smaller only one fourth or one sixth of this size. When wetted, the grains of pollen promptly imbibe walcr l>y endosmosis. and are distended, changing their shape somewhat, and obliterating the longitudinal folds, one or more in number, which many grains exhibit in the dry slate. Soon the more extensible and elastic inner coat inclines to force its way through the weaker parts of the outer, especially at one or more thin points or pores; sometimes forming projections, when the absorption is slow and the exterior coatiii<>- loii-di. In O S many kinds of pollen, the grains, when immersed in water, soon distend to bursting, discharging the contents.1 474. Pollen-tubes. In others, and in most fre>h pollen, when placed in ordinarily aerated water, at least when this is slightly thickened by syrup or the like, and submitted to a congenial tem- perature, a projection of the inner coat through the outer appears at some one point, and by a kind of germination grows into a slender tube, which may even attain two or three hundred times the diameter of the grain ; and the richer protoplasmic contents tend to accumulate at the farther and somewhat enlarging ex- tremity of this pollen-tube. - 475. In cleistogamons ilowers (l.'il). the pollen, while still in the anther, sends out its tubes, which may grow to a great length, in the mere moisture of the llower-bud. the growing tip always directing itself toward the stigma in a wonderful way. Similarly, in the open flower of Milkweeds, the pollen-tubes sometimes start from the pollen-mass even while yet in the anther, and in vast numbers, forming a tuft or skein of pollen-tubes, which may attain considerable length and direct itself toward the some- what distant stigma. Commonly, however, the pollen remains 1 In Con i ferae, the grains of pollen have a peculiar internal structure or nil her a development ( surest ive of a homology with the microspores of some of the higher ( 'rypto^amia ), tin- contents at or before maturity iiinlfr^oin^ ilivision into two or three internal cells, only one of which acts in fertiliza- tion When they act upon the ovule or ;m> placed in water, and the inner coat swells by absorption, tin- hurstinij outer coat is commonly thrown off. In I'ines and Firs (hut not in Larch and Hemlock Sprue.-), the grain of pollen is singularly compound, consisting (as in Kiir. f>]S| of a central arcuate liody (the proper pollen-cell) hearing at each end an empty roundish cell. The.-.- are vesicular protrusions or appendages of the proper pollen-irrain, of no known functional importance, except that they render such wind-dis- persed pollen more buoyant for transportation. 2 Van Tie^hem, in Ann. Sci. Nat. ser. 5, xii. 312, &c., 1869. GYNCECIUM IN ANGIOSPERMS. 259 unaltered until it is placed upon the stigma. The more or less viscid moisture of this incites a sim- ilar growth, and also doubtless nour- ishes it ; and the protruding tube at once penetrates the stigma, and by glid- ing between its loose cells buries itself in the tissue of the style, descending thence to the interior of the ovary and at length to the ovules. Fertilization is accomplished by the action of this pollen-tube upon the ovule, and upon a special formation within it. Consequent upon this an embryo is formed ; and the ovule now becomes a seed. SECTION VII. THE PISTILS, OR GYNCECIUM. § 1. IN ANGIOSPERMS. 476. The succinct description of the pistil in the first section of this chapter (302), as also what has been stated of the modi- fications of the g3'noeciuHi in Section III., relates to the most typical conditions of this part of the flower. The essential characteristics of all ordinary pistils, whether simple or compound, are : 1 . a closed ovary, in which one or more ovules are included ; and 2. a stigma, upon which pollen for fertilizing the ovules is received, and through which the pollen acts upon them. There is a more simplified condition, in Gymnosperms, in which naked ovules are exposed to the direct action of the pollen. In con- tradistinction to this, the ordinary pistil is said to be Angiosper- mous ; that is, with the seeds enclosed in a sac or covering, this in the flower being the ovary.1 And plants with such gyncBcium are denominated ANGIOSPERMS or ANGIOSPERMOUS PLANTS. To such only the present subsection specifically relates. 477. The several terms which apply to the Gyncecium or female system of a flower, and to its components, have been 1 Although thus originated, the seeds are not in all cases matured in a closed pistil. In the Blue Cohosh, Caulophyllum thalictroides, the ovules rupture the ovary soon after flowering, and the seeds become naked ; and in Mignonette they are imperfectly enclosed, the ovary being open at the summit from an early period of fructification. FIG. 523. A pollen-grain of Datura Stramonium, emitting its tube. 524. Pollen- grain of a Convolvulus, with its tube. 525. Other pollen-grains, with their tubes, less strongly magnified. 526. A pollen-grain of the Evening Primrose, resting: on a portion of the stigma, into which the tube emitted from one of the angles penetrates; the oppo- site angle also emitting a pollen-tube. All highly magnified. THE FLOWER. enumerated and defined already (302, note) : the elementary term is that of 478. Carpel, Lat. C'AKPI.I.I.IM. This is the term coined by Dunal, and is in common use. The better-formed word CAKI'IIUTM (English ('(ir/nd) lias been proposed, and best of all CAKPO- I'liYi.i.r.M, in Fnglish Carpopliyll. For carpels are. as the word carpophylla denotes, pistil-leaves, or leaves of the gymrcium, /. i'., seed-bearing or fruetiferous phylla. The}- occupy the cen- tral or uppermost region of the flower. A carpel may IK- a pistil of itself, either the only one of a blossom or one of several, or it may be a constituent of a more complex pistil. In either case, a carpel is the homologuc of a leaf. 17'.). The morphological conception of an uncombined carpel is that of the bhule of a leal' incurved lengthwise, so that the margins meet, and join by a suture, thus forming a closed sac, the ovary. A prolongation of the tip of the leaf is the sti/fr : some portion of this, usually the apex, not rarely a single or double line down the side which answers to the suture of the leal-margins, and may be regarded as its continuation, is the stigma. The carpellary leaf is alwaj-s incurved : the lower sur- face of the leaf is represented by the exterior surface of the ovary, the upper by the interior. The conjoined margins of the leaf, or whatever they bear, are internal in the ovary: the stigma may lie regarded as a portion of leaf- margins presented externally, des- titute of epidermis and formed of loose cellular tissue, which in anthesis is moist by some secretion. The ovules are peculiar structures normally arising as outgrowths from the margins of the leal', or some part of them, sometimes from the whole or a special portion of the upper or inner surface of the leaf. •LSI). The carpellary leaf being involute, the suture, on which the ovules are normally borne, always looks toward the axis or <-eiitre of the Mower. It is the only proper suture (or seam) a carpel can have. From its position it lakes the- name of Inner or }'<'ntr."> 1 divides the cell of a singn- carpel. Such are found in Flax (Fig. f>3!>-.Vl 1 ) . in Amelanchier or Service-berry, in Huckleberry (< layhissaeia) . and in most of FIG. 535. T'istil of :i S;i\ilY:i-< nipnsr.l ,.r two carpels or simple pistils united below l.ut distinrt abOTe; cm acrOBB l»>th al.ove :nul below. FKi. r,:;6. Pistil of common St. Johnswort, of three united ovaries; their distinct. FK!. 5.'!7 The same of another species <.f St. -Tohnswort (Hvpencum prolificum), tin- Mvlrs ,-ilso niiili'il into on.>. whi.-h. however, limy split ;ip:irt in tin- fruit. FIG. 53S. I'i-iil of 'I'T-.-Llrsrantia or Si.i.l.'rwort. even the three sti-nias united into one. The ovary in all cut across to show the internal structure GYNCECIUM IN ANGIOSPERMS. 265 the American species of Vaccinium. In all these, the false par- tition is a growth from the middle of the back of each carpel, which divides its cell more or less completely into two. 540 541 494. On the other hand, even the true dissepiments which belong to such a compound ovary may be abortive or evanescent, the placentae remaining in the axis combined into a column. (499.) The second modification of the compound pistil (491) normally has an ovarj^, 495. With one Cell and Parietal Placentae. That is, the placentae are borne (as the term denotes) on the wall or parietes of the ovary, as in the Poppy, Violet, Sundew, Cistus or Helianthemum (Fig. 543), Cleome, Gen- tian, and in all or most of the orders from which these examples are cited. The diagram Fig. 542 illustrates the morphological conception of a com- pound pistil of this kind. Not that it is ever sup- posed to be formed by the actual combination of once 544 545 separate leaves, any more than a gamophyllous calyx or corolla is actually so produced. The conception in all such cases is that FIG. 539. Transverse diagrammatic section of a flower of the common Flax, show- ing the ovary with false partitions extending one from the back of each cell. 540. Sec- tion of a mature fruit and seeds of the same, the false partitions now complete, divid- ing the five cells into ten. each one-seeded. 541. Same of a wild Flax (Linum perenne), in which the false partitions remain incomplete. FIG. 542. Plan of a one-celled ovary with three parietal placentae, cut across be- low; the upper part showing the top of the three leaves it is theoretically composed of, approaching, but not united. FIG. 543. Ovary of Helianthemum Canadense, cut across, showing the ovules on three parietal placentae. FIG. 544. Transverse section of the ovary of Hypericnm sraveolens; the three large placenta; meeting in the centre, but not cohering. 545. Similar section of a ripe capsule of the same; the placentae now evidently parietal 266 THE FLOWER. of a congenital development of organs in union which, in the development of a vegetative shoot, would lie leaves. This case is represented by the combination of open carpellaiy leaves, as the preceding one is by that of closed ones. As the edges of the leaves must needs be turned in, to bear the ovules, a compound ovary with parietal placentation may be likened to the unopened calyx of a Clematis, as shown in Fig. iVii;. 257. Every gradation is found between axilc and parietal placentation. Sometimes the placent;e are strict!}- on the pari- etes or wall (Fig. 543, 5-17) ; sometimes borne inwards on incomplete dissepiments (Fig. 5-1 ;>()) ;1nd Portnlacacere. this evidently results from the obliteration of the dissepiments (as many as there are styles or stigmas), vestiges of which may be sometimes FIG. f>4f> l>i;iL.'r:iMi (u'ri>und-pl:iiO to illustrate froc i-i-ninil i>l.ioentation produced by abortion of dissepiments. 547. Same of strict parietal placentation. 548. Same with the placenta carried inward on imperfect dissepiments. GYNCECIUM IN ANGIOSPERMS. 267 detected, while certain plants of the same families, of otherwise identical structure, retain the dissepiments even in the fruit. 500. But a similar condition may equally arise from a modification of parietal placentation, namely, with the margins of the leaves ovuliferous only at bottom, and the placentae there conspicuously devel- oped and completely united. The basal placenta- tion of Dionaea is unavoidably so explained, its nearest relative, Drosera (Fig. 553) , having parietal placentae. And this leads to a probable explanation of the case in Primulacea?, where a large free central placenta fills the centre of the cell, and no trace of dissepiments can be detected.1 501. The idea maintained in former editions is still adhered to ; namely, that placenta? belong to carpels and not to the cauline axis, in other words, that ovules are productions of and borne upon leaves, usually upon their margins, not very rarely upon other portions of their upper surface, rarely- over the whole of it.'2 502. Ovules cover the whole internal face of the carpels in Butomus and its relatives, also of the AVater-Lilies (both N 3-111 phffia and Nuphar, Fig. 551) excepting the inner angle, to whiclr the}' are usually restricted in other plants. And in the allied Brasenia and Cabomba, where the ovules are reduced to two or three, one or more of them is on the midrib, but none on the 1 The placenta in this and like cases is rather to be Regarded as an out- growth from the base of the carpellary leaves, combined over the floral axis. Upon this interpretation, a central portion of the column may be (and sometimes must be) of axile nature, yet the ovules be borne upon foliar parts. See Van Tieghem, in Ann. Sci. Nat. ser. 5, xii. 329 (1809) ; Celakowsky, Vergleichende Darstellung der Placenten, &c. (1876) ; Warming, in Ann. Sci. Nat. ser. 6, v. 192. 2 This view was first maintained as a general theory, and on critical grounds, by Brown, in Plants Javanica Rariores, 107-112. Schleiclen, End- licher, and others took the opposite view, /. e., that ovules are productions of the axis, even in parietal placentation, — an exceedingly far-fetched suppo- sition. In later days, the commoner view has regarded ovules as of both origins, as productions of the carpels in parietal, of the axis in at least some free central or basilar placentation. But at present the theory of foliar origin without exception, revindicated by Van Tieghem, and espe- cially by Celakowsky and Warming, again prevails. For the bibliography and an abstract of the various views, see Eichler, Bliithendiagramme, espe- cially the note in the preface to the second part (where be gives his entire adhesion to this conclusion) ; also Warming's memoir, De 1'Ovule, in Ann. Sci. Nat. ser. 6, v. 1877-78. FIG. 549. Vertical section through the compound tricarpellary ovary of Spergularia rubra, showing the free central placenta. 550. Transverse section of the same. THE FLOWER. margins of the carpellary leaf. In many species of Gentian, as :ilso in Obolaria and Bartonia. of tin- same family, the whole internal face of a dic'ii- pellary ovary is thickly ovulifermis. .")!):;. Perhaps lln- parietal placenta- in Parnassia (Fig. 552) are borne <>n the midribs of the carpels, for they are directly under the stigmas, instead of alternate with them, as they normally should be. The same thing occurs in Poppies and many other Papaveraceae, also in some Cruci ferae ; and in some of the cases each stigma is more or less two-lobed. This sug- gests the explanation,1 here probably the true one. which supposes that the placentae are borne on the leaf-margins in the normal way, but that each stigma is two-parted (as if the carpel- lary leaf were deepby notched at the apex, and so its two stigmatic leaf- margins separate, as Drosera illus- trates. Fig. 5.">;!). and th:,t the two half-stigmas of adjacent carpels have coalesced into one body, which would of course stand over the parietal placenta- beneath. Each stigma in such a case, as well as each parietal placenta, would consist of the united margins of two adjacent carpels. 553 § 2. IN GYMNOSPERMS. 504. GYMNOSPERMOUS (that is, naked-seeded) plants are so named because the ovules, or bodies which are to become seeds, are fertilized by direct application of the pollen, which reaches and acts upon the nucleus of the ovule itself, not through the mediation of stigma and style. In the structure of their (lowers. these plants are of a low or simplified type, in some respect > nol obviously homologous with the Angiosperms which now consti- tute the immense majority of pha-noganious plants. But, up to a comparatively late geological period. ( iymnosperms appeal' to have been the only flower-bearing plants. They are represented 1 Given by Brown, in tin- I'liinta- Javanica> Kariorcs, above ri'JVrml to. FIG. 551. Transverse seet ion of ;ui ovary of Nyruphiea odorata, the carpels ovulifer- OUS over the whole interior surface. FIG. 552. Pistil of I':n nassi.-i. with ovary transversely divided. FIG. 553. Pistil of l>n>sera liliformis, with ovary transversely divided. GYNCECIUM OF GYMNOSPERMS. 269 in the extant vegetable kingdom by three (or four) groups or orders, two of them small, and one comparatively ample and of wide distribution ; and all are so strikingly different from each other that the}- cannot be illustrated by a common description. The largest order, Coniferae, is familiar, and contains a good share of the most important forest trees of temperate climates. The smallest, Gnetaceae, chiefly tropical or of warm regions, lies between Gymnosperms and common Dicotyledons. The third, Cycadaceas, is most remote from them, and as much so from Monocotyledons, except that it imitates Palms, as it also does the Tree-Ferns, in habit, both as to stem and foli- age. The particular morphology of Gymnosperms would re- quire for its illustration copious details and the history of various conflicting hypotheses. It must be relegated to the special morphology of the natural orders, premising, however, a brief sketch of the general floral structure.1 505. In Gnetaceae, Gymnosperms and Angiosperms almost come together. The flowers have a perianth (diphyllous or tetraphyllous) ; the stamens have a distinct filament and anther ; and the gynoecium is a sac (presumably of two carpophylls) open at the top and filled at bottom by a single ovule of the simplest kind, i. e. consisting of a nucleus destitute of coats. This pistillary body is attenuated and prolonged above the ovule into a style-shaped tube, with open and commonly two-cleft orifice. In the almost hermaphrodite sterile flower of Welwitschia, this takes the form of a much dilated stigma, which is even beset with seeming stigmatic papillae. If only the pollen were here to grow forth into pollen-tubes (with or without a closing of the tube) , angiospermy would be attained. But, in fact, the pollen- grains bodily reach the ovule itself through the tube, fertilizing it directly.'2 This interesting group of plants consists of the 1 References to the literature of gymnospermy and to the steps of the prolonged controversy over it, also the points of morphology still in part unsettled, need not here be given. The history and the idea of gymnospermy began with Robert Brown's paper on Kingia, " witli Observations .... on the Female Flower of Cycadeas and Coniferas," read before the Linnean Society in the year 1825, and published in King's Voyage in 1827 ; and the bibliography down to a recent date is given by Eichler in Flora Brasiliensis, Gymnospermia, iv. 435, and in Bliithendiagramme, i. 55-69; also ii. preface x. See also Alph. DeCandolle, Prodr. xvi.- 345, 524. In this volume, the late Prof. Parlatore adhered to the ancient ideas in his monograph of the Coniferaj. 2 The view here implicitly adopted is that of Beccari, founded on the study of Gnetum, and published in Nuovo Giornale Botanico Italiano, ix. 1877. It was before nearly or quite reached in successive steps, by J. D. Hooker, in his classical memoir on Welwitschia, in Trans. Linn. Soc. xxiv. ; Stras- burger, Die Coniferen und die Gnetaceen, 1872; and "W. R. McNab, in Trans. Linn. Soc. xxviii. 1872. 270 THE K LOWER. genus dnetiim, shrubs or trees, with nearly the aspect of Angiosperms, having broad and pinnately-veined leaves : \Vel- witschia of tropical AY. Africa, remarkable for its persistent cotyledons which form the only foliage of a wood}' and long- eiiiluring plant, and for its stem or trunk which broadens with- out lengthening, except in its flower-stalks : also Kphcdra, of much branched shrubs, mainly of warm-temperate regions, leafless or nearly so, one species of which inhabits Europe and two the southern borders of the 1'nited States. 50G. The flowers in all Gyninospeniis are diclinous, either dioecious or monoecious ; except that those of the si range < ineta- ceous genus \Yel\vitsehia are structurally polygamous, the male flowers having a well-formed but sterile gymecium. 507. In Conifera', the largest and most important tvpe, are embraced all the familiar (iymnospcrms of temperate regions, Pines, Firs, Cedars, Cypresses, which bear their flowers in catkin-like clusiei-s and their fruit in cones, and also the Yews and allied trees which do not'' produce cones. Perianth being want- ing and the sexes wholly separate, the floral type is so degraded that it becomes doubtful whether cadi cluster of anthers, or of ovnlil'erous scales or ovules, constitutes a blossom or an inflores- cence. Certain botanists look upon a whole catkin, and others upon a male catkin only, of a Pine or Fir as forming one llower. It is here assumed that each stamen of the one and each ovu- liferous scale of the other answers to a flower of the simplest sort. ' The anthers are extrorse, the cells or pollen-sacs b<>longing to the outer or lower side of a scale or a 1 It will lie seen that, for the female (lowers, tins follows of course from generally accepted view ; and, where tin's is conceded, analogy may extend it to the male catkins also: yet in such cases, where all thephylla of an indefinite simple axis are stamens, spirally arranged on it, the difference hetween inflorescence and male Bower completely vanishes. Kit!. r>~>-\. Female llower .it' a Yew. an ovule surrounded l>y its liracts. 555. Longi- tudinal nnil moiv enlarged section of a female tlower of Yew ami of the upper part of the si 101 it it. terminates : I lie thick coat of the ovule open at t hi' top, the nucleus within, anil thccaisnc. 657 GYNCECITJM OF GYMNOSPERMS. 271 559 connective : sometimes these sacs or cells are two, and the organ evidently homologous with an ordinary stamen : often they are more numerous (from three to twent}*) and variously disposed. 508. The Yew Fam- ily (Taxineae) is next to Gnetaceae in structure. It is generally ranked as a suborder of Coniferae, but it may claim to be a distinct order. The gyncecium is a naked ovule, terminating a stem,1 and surrounded by several bracts. After fertilization, an outgrowth of the receptacle (or a kind of disk, 394) makes its appearance as a ring girding its base : this grows in height and thickness, and becomes a soft-fleshy cup, imitating a hollow berry, in the bottom of which the stony-coated seed nestles. (Fig. 554-557.) Very similar is the gyncecium of Torreya, except that the cup- shaped disk develops almost simultaneously with the ovules, and as it grows becomes adnate to the large seed in the form of a fleshy coating. In the Gingko, two or more similar ovules are nakedly developed on a naked peduncle, un- accompanied even by a bract (Fig. 558) , and one or more of these ripens into the berry-like seed, Fig. 559. In Podocarpus there are some sub- tending bracts, and the naked ovule 1 It does not therefore follow that the ovule is a part of the axis, or is terminal in the sense of being its direct continuation. In this regard it may be only what the pistil of a Cherry is, which to all appearance is equally a terminal production, but is really the representative of the last leaf of the axis. If so, that leaf is here suppressed to the utmost, and replaced by what is ordinarily its outgrowth, the ovular nucleus and its coat. The structure of Podocarpus favors this interpretation. FIG. 558. Female flowers of Gingko biloba or Salisburia adiantifolia. 558«. Portion of the same enlarged. After Strasburger. 559. A drupaceous seed of the same, in vertical section, exhibiting the mature disk which forms the flesh, the crustaceous seed- coat, within which is the kernel of the seed; at the base on one side a sterile ovule is seen. After Decnisne. FIG. 560. Female flower of Podocarpus (an ovule inverted on a column or elevated support), subtended by bracts. After Eichler. FIG. 561. Magnified vertical section of a similar flower of Podocarpus. After Strasburger. 272 THE FLOWER. is inverted on a more or less lengthened and stout support, which is conceived to represent the carpel. (Fig. 5GO, 561.) r>een much controversy over the morphology of the parts. With the former, the discussion turns on the character of the ovulil'cnuis scale. As to this, the hypothesis originally proposed hy Mohl, and adopted by Braun, is now said to he satisfactorily demonstrated hv Sten/el. in Nov. Act. Nat. Cur. xxxviii. 1S7G. See note hy Kngclmnnn in Amer. .Tour. Rci. Dec. 1876, and also the preface to the second part of Eichler's Bliithciidiugramme, Fl< ; -,«;•_'. View of the upper face of a carpellary scale of a Lan-h, showing the pair of adnatr rid. 563. Similar vi.-w ..f a rarpollary scale of :i Larrh. and of a l>ract behind it. 564. Ground plan <>f tin- satin- in diagram, reversed; the upprr ligmc di -not ing the axis of the cone, the lower tin- In-net, the middle one the carpellary si-ale and the two ovules borne on its face. After Eichler. GYNGECIUM OF GYMNOSPERMS. 273 511. In the Araucaria tribe the ovuliferous or carpel-scale is throughout smaller than the bract, and is completely adnate to it, or with only the tip free ; that of Araucaria (Fig. 565) bears only one ovule, high on the carpel, the orifice downward as in the Pine tribe. In Taxodium, Sequoia, and the like, the cone- scale is equally inferred to be composed of bract and carpel-scale united ; and indications of this composition are to be observed. The ovules (from two to several) are at the base of the scale, erect and free. The cone-scales are alternate and spiral on the axis, but indistinctly so in Taxodium, the Bald C}rpress or so-called Cypress of the Southern United States. 512. In the true Cypress tribe (Cupressinese) the cone-scales, which are never numerous, are opposite or verticillate, i. e. like the foliage- leaves, in whorls of twos, threes, or sometimes fours ; and the ovules are from two to 568 1878, where it is fully adopted. It was suggested by certain rather common monstrosities, and by the two combined leaves of Sciadopitys. According to this view, the ovuliferous scale in the Pine tribe is com- posed of two leaves of an arrested and transformed branch from the axil of the bract, which are in the normal manner transverse to the subtending bract, are here carpellary, each bearing an ovule on the dorsal face ; the two are coalescent into one by the union of their posterior edges, and the scale thus formed is thus developed with dorsal face presented to the axis of the cone, the ventral to the bract. It is therefore a compound open carpel, composed of two carpophylls. This character of being fructiferous on the back or lower side of the leaf occurs in no other phsenogamous plants, but is the rule in Ferns, from something like which Conifers may be supposed to have been derived ; the ovules of the one in this regard corresponding to the sporangia of the other. FIG. 565. Vertical section (in diagram) of a bract, adnate" carpel-scale, and adnate ovule of Araueari.a imbricata. After Eichler. FIG. 566. Bninuhlet of the American Arbor- Vitse, considerably larger than in na- ture, with a forming fertile cone 567. One of the scales removed and more enlarged, the inside exposed to view, showing a pair of naked erect ovules on its base. FIG. 568. Fertile flowers of true Cypress (Cupressus sempervirens), after Baillon: a forming cone, with one scale cut away, to show the cluster of ovules under it. 18 L'74 THE FLOWER. several at or on the base of each cone-scale, always with orifice upward. Arbor-Vita' (Fig. 5GG, 5G7) has a single pair of ovules to the scale ; Junipers, sometimes only one ; true Cypresses (as in Fig. 5G£), often a dozen or more. At flowering time, the cone- scales mostly appe:ir as if simple ; but in most genera they soon thicken greatly within ; and they are usually understood to be composed of bract and carpel-scale combined, the latter of the same constitution as that of Pines and Spruces, but perfectly consolidated and confluent with the bract-scale.1 S7:> 513. In Cycadacese, the type of the flower of Angiosperms is almost or quite lost; yet the organs maybe h.omoloui/.ed with those of Coniferoe, which these plants are wholly unlike in habit. 1 This internal and ovuliferous scale may seem to be wholly hypotheti- cal, and assumed to homologize the cuprcssineous with the abietineous cone. Without it, \ve should have to consider that, while in Abictineaj the ovules belong to leaves of a secondary axis, in fupressinea' thev are borne on those of a primary axis, or else are axillary productions without carpels. But in the Arauearia tribe the internal scale is obvious ; and there are suffi- FIG. 5G9-r>7.ri. /amia, diielly 7.. media, after Riolmrd. fifi!). A main plant. 570 Lower part of a male catkin. 571. A stamen removed, showing inimeron- small pollen- sacs under the peltate top. 572. A female catkin, with a quarter section ent away. 573. A female flower or carpel, with two enlarging ovules or yoiinvr seeds. ."74. Ripe seed, with the thick fleshy coat cut away at apex. 575. Longitudinal section of ripe seed, more enlarged. GYNCECIUM OF GYMNOSPERMS. 275 Their likeness to Palms and other Monocotyledons is confined to the port of their unbranched trunks and their pinnate leaves with parallel-veined or simple-veined leaflets ; nor have the}7 any further resemblance to Ferns, except that in some the leaflets are circmate in vernation. Although a tropical type (of small present importance, compared with the part which it played in the Devonian and Cretaceous periods), it has one small representa- tive (Zamia media, the Coontie) at the south-eastern extremity of the United States, and a more striking one (C}7cas revoluta, well known in cultivation) in the southern parts of Japan. 514. Following the analogy of Coniferae, each scale (whether of the pollen-bearing or the ovule- and seed-bearing ament) of Zamia (Fig. 569-575) is here regarded as a flower. Here the phylla, or scales with peltate top and stalk-like base, are exter- cient indications of similar composition in the cupressineous cone-scales tc induce the adoption of it by Parlatore, who rejected the idea of gymnospermy ; and, finally, this composition is nearly demonstrated by VanTieghem (1868) upon the anatomical structure, and by Strassburger (1872) on the development. FIG. 576-578. Carpophylla of Cycas revoluta, much reduced in size. 576. One bearing ovules below and leaflets or leaf-lobes towards the apex. 577. A similar carpo- phyll with leaf-lobes reduced to mere teeth, and ovules in place of the lower teeth. 578. A similar carpophyll in mature fructification, bearing the large drupaceous naked seeds. The last two after Richard. 276 THE FLOWER. nail}- much alike in the two sexes, which throughout the family occupy separate plants. The male flower (Fig. 068) or .stamen. if it may he so termcil. bears indefinite pollen-sacs on the under side of the peltate portion, sometimes extending to the upper part of its stalk. The homologous female flower, or carpophyll, bears a suspended ovule on each side of the stalk (Tig. 573), which becomes a large fleshy-coated seed. In Cycas the male ament is not very dissimilar, although on a larger scale. lint the earpophylls are evident leaves, not condensed into an ament, but loose or spreading, of a character and aspect intermediate between the lax bud->cale> which precede and the pinnate foliage- leaves which follow them in development. Along the margin of what would be leal-blade they bear ovules in place of leaflets, lobes, or teeth (Fig. 57G-578) ; and these, when fertilixed from the male flowers, mature into large and drupaceous naked seeds. Even without fertilization, such seeds grow to their full size on the lemak' plant of the common Cycas (or falsely so-called Sago Palm) , but form no embryo. SECTION VIII. THE OVULE.* 515. Ovules (302) are peculiar outgrowths or productions of carpels which, upon the formation of an embryo within, become seeds. In the angiospermous gyncecium (470) they are nor- mally produced along the margins, or some part of the margins. of the carpellary leaf (-ITS), either immediately, or by the in- termediation of a placenta ( IN.")), which is a more or less evident development of the leaf-margins for the support of the ovules. Rarely, yet in a considerable number of cases (501, 502). ovules are developed from the whole internal surface of tin- ovary, or from various parts of it. in no definite order, directly from the walls, and without the intervention of any thing which can be regarded as placenta. In ( lymnosperms (501-514) the ovules are borne on the face of the earpellary scale or at its base; or on leaf-margins, as in Cycas; or, when there is no representa- tive of the carpel, on the cauline axis, seemingly as a direct growth of it. (508, note.) 5HJ. As to attachment, ovules are either sessile, i. e. stalk- less, or on a stalk of their own (Fig. 582, 584), the FUNICULUS or PODOSPEKM. As to number they are either solitary, few, or 1 Lat. Ovulum, pi. Ovula, diminutive of ovum (egg), perhaps first used by Adanson OVULES. 277 indefinite \y numerous. They may also be indefinite or variable in number when not particularly numerous. 517. As to situation and direction within the ovary, the terms are somewhat special. Ovules are erect, when they rise from the very bottom of the cell, as in Fig. 580 ; ascending, when attached above its bottom and directed upward, as in Fig. 579 ; horizontal, when borne on one or more sides of the cell and not directed either upward or down- ward, as in Fig. 314, 315, 530; pendulous, when more or less hanging or declining from the side of the cell ; suspended, when hanging from the apex of the cell, us in Fig. 581. 518. The body and only essential part of an ovule is its NUCLEUS. This in most cases is invested by one or two proper coats. The coats are sacs with a narrow orifice, the FORAMEN. In the seed, the closed vestige of this orifice is termed the Micropyle ; wherefore this name is sometimes applied to it in the ovule likewise. When the ovule has two coats, the foramen of the outer one is called EXOSTOME, of the inner ENDOSTOME ; literally the outer and the inner orifice. The coats themselves have been named PRIMINE and SECUN- DINE, but with an ambiguity in the application which renders these names unadvisable : for in their formation the coats appear later than the nucleus, the inner coat earlier than the outer ; and the name of primine has by some writers been applied to the earlier formed, by others to the external coat. The proper base of the ovule, from which the coats originate and where these and the nucleus are confluent, is the CHALAZA. The attachment of the ovule to its funiculus or support, which in the seed becomes the FIG. 579. Ovary of a Buttercup, divided lengthwise, tn display its ascending1 ovule. 580. Same of Buckwheat, with an erect ovule. 581. Same of Anemone, with a sus- pended ovule. FIG. 582. Diagrammatic section of a typical or orthotropous ovule (such as that of Fig. 582"), showing the outer coat, a, the inner, b. the nucleus, c . the chalaza. or place of junction of these parts, rf. (The coats are never so separated and the nucleus so re- duced in size as is represented in this mere diagram.) 583. An ovule similar to the preceding, but curved, or campylotropous. 584. Au amphitropous ovule. THE FLOWER. HILUM, takes also this latter name in the ovule. In the simplest form of ovule (as in Fig. 5*2, 5*0). hiluin and ohala/a are one. So also in cases where the body <>!' the ovule- incnn e>. as in Fig. 583. But verv commonly tlic place of attaclinifiit. which becomes the hiluin, is more or le.ss distant from the clialaxa ; as in Fig. ~>*\ and .")*?, where the hiluin i,- lateral, but the diala/a at the larger end. the two being connected l>y a short ridge : and in Fig. 588 the two are separated by the whole length of the ovule. 519. The simplest and most rudimentary ovule is that with- out a coat, as in Mistletoe and the whole order Loranthaeea1, and in Santalaceae and (inetaceae. This has been called a naked ovule; but long before ovules of such simplicity were known this term had been appropriated to those of Gymnosperms, in the sense of destitute of ovarial or pericarpial covering, i. e. to uncovered ovule, not to uncovered nucleus. The ovule consist- ing only of nucleus may be termed (after Alph. DeCandolle) simple, or better achlamydeous.1 520. The tunicated or chlamydeous ovule is of three principal kinds, with one or two subordinate modifications. These are the orthotropous, campylotropous, and anatropous, and the modi- fication called half-anatropous, or amphitropous.2 521. Orthotropous (Fig. 580, 582, 585), or straight ovule, is the simplest but least common species, being that in which the 585 chalaxa is at the evident base, and the oriiice at the opposite extremity, the whole ovule straight (as the first part of the name denotes) and svmmetrical. Afropous, meaning not turned at all, is a later and et3'mologically much better name, but it has 1 An epidermal stratum or tegument may not be wanting to such ovules, forming a sort of adherent rove-ring; but this in nature and origin is not similar to the uvular coats. 2 In Latin form. tii-tlmim/Hi. campylotropa, tnitit/-<>/>iii>n.s nr AtrOpOOS OTUlO Of Baofewbeat. r.si;. ovule of rliiokiv, ,l. r,S7. Vnipliitropnua OTUle of Hallow, fiss. Anntrnpnus ovule of a Violet. Tlie letter /( indicates the liilum; r, the clmlaza. \vhich in 585 and 580 corre- sponds to the liilum; /, the foramen or orifice; r, the rbaplie. OVULES. 279 not come into general use. This ovule is characteristic of Polygonaceoa, the proper Urticaceae, Cistaceae, &c. 522. Campylotropous (Fig. 583, 58G) is the name of the ovule which in the course of its growth is curved on itself so as to bring the orifice or true apex down close to the base, here both chalaza and hilum. This and the orthotropous ovule begin their development on the placenta in the same way, but the camp3'lotropous develops unequally, one side enlarging much more than the other, especialty at the base, until the ovule becomes reniform, and chalaza and orifice are brought into O close proximity. Campylotropous ovules are characteristic of Cruciferae, Capparidaceae, Resedaceae, Caryophyllacese, and Chenopodiaceae. 523. Ainphitropons (Fig. 584, 587), also termed Heterotropous and sometimes Half -anatropous, is between the preceding and the following ; and it passes in various instances either into the one or into the other. The body of the ovule is straight or straightish, but it stands as it were transversely or at right angles to the funiculus and hilum ; and it is fixed ~by the middle, the chalaza at one end, the orifice at the other. An apparent continuation of the funiculus, adherent to the outer coat, extends from the hilnrn to the chalaza. Compared with the preceding form, the explanation is, that the unequal development at its formation is confined to the basal half, and the axis remains straight, while the whole is half inverted by the very unequal growth. Compared with the next form, the inversion is less and the later growth or extension of the apical portion greater. The amphitropous ovule is characteristic of Primulaceae, and is common in Leguminosae. 524. Anatropous (Fig. 588, also 579, 581, 597) is the name of far the commonest species of ovule, that in which the organ, under the course of its growth, is quite inverted on its base ; so that, instead of standing at right angles with the funiculus, it is parallel with it, or rather with the apparent continuation of it, which is adherent to its surface as a sort of ridge or cord extending along the whole length of the ovule, from hilum to chalaza. The latter occupies the seeming apex of the seed ; and the organic apex or orifice is at the other end, close beside the hilum. At maturity, the ovule is straight, but not wholly symmetrical, the attachment being oblique or somewhat lateral, and the ridge or cord on that side not rarely prominent. 525. The cord or ridge, which extends along the whole length of the anatropous ovule, and for half its length in the amphi- tropous (Fig. 588, 587, r), is named the RHAPHE. This is not 280 THK 689 at all a scam, as the Greek word denotes. Its origin, and the whole structure of such ovules will be apprehended by comparing various stages of its growth. 526. An ovule of any kind at the beginning is an excrescence or outgrowth of the placenta, or of some part of the leaf-surface if there is no developed placenta. This incipient ovule is the nucleus (518), or the nucleus surmounting a rudimentary funiculus. The nucleus is soft cellular tissue only, from first to ':ls^- The achlamydeous ovule (519) undergoes no further development except in size or shape. Indeed sometimes (as iu Balanophorese) this bare nucleus is reduced to a few cells of parenchyma. 527. In ordinary ovules a new growth early begins around the base of the nucleus, or is sometimes coetaneous with it, at first as a ring (or part of a ring), soon as a cup, at length as an enclosing sac or covering, open at the top ; this is the inner coat of the ovule when there are two. The outer coat begins and goes on in the same way, and at length grows over and encloses the inner coat as that did the nucleus. (Fig- 590-595.) When- ever there is a third and more exterior coat it is formed during the growth of the fertilized ovule into the seed, to which there- fore it belongs, and in which it takes the name of arillus. (597.) At the time of fertilization the apex of the nucleus, or a pro- longation of it, usually projects beyond the orifice and there receives the descending pollen-tube. Some fibre-vascular tissue, especially spiral ducts, may be found in the funiculus and cha- la/a, sometimes extending into the coats. 52S. The development of the orthotropous or atropous (un- turned) ovule proceeds symmetrically, without distortion, the parts keeping their primitive direction. In the campy lotropous, the whole of one side of the ovule greatly outgrows the other. !'!<;. M'.i. Magnified view of a vertical section of a carpel of Magnolia Umbrella, about a month lieibre anthesis, showing one of the two nascent ovules, at. this time only nucleus. FIG. 590-597. Further development of the ovule of Magnolia I'mln-ella. sin. wing the formation of the e.oats and the anatropy. 500. Ovule a week older than in 58!». 691. Same a week or two later. 502. Same :\ few .lays later r.o:! Same from a nearly full- grown flower-bud. r,!U. Same at timeof authesis. 595. Vertical section of the last through the middle of the rhaphe. 596. Cross-section of the same. (See Jour. Linn. Soc. ii. 108.) OVULES. 281 In the anatropous, the inequality of growth is mainly confined to the base or chalazal region, which ends by becoming upper- most ; and the full-grown ovule has the ap- pearance of being inverted on and adherent to the upper portion of its funiculus, the rhaphe. Fig. 589-597 illustrate the course d of development from a comparatively early period. 529. The direction of anatropy or of other turning of the ovule in the course of growth is somewhat diverse. But in general, when- ever ovules are in pairs, the two turn from 597 each other, in the manner of Fig. 315, and so present their rhaphes back to back. The rhaphe-bearing may therefore be called the dorsal side of the anatropous ovule. The same is true in the case of numerous ovules, viz., those of one half of the placenta (or one leaf-margin) turn their backs to those of the other. When such ovules are solitary or in single rows, and either ascending or hanging, the rhaphe is usually on the side next to the placenta or ventral suture, as in Fig. 579 : it is then said to be ventral (i. e., next the ventral suture), or adverse to the placenta. In certain cases, mostly in hanging ovules, as in Fig. 581, the rhaphe looks in the opposite direction, toward the dorsal suture or midrib of a simple ovary : it is then said to be dorsal or averse from the placenta.1 «J 1 By comparison of Fig. 578 with 576 and the like, it may be perceived that the difference is explicable oy a kind of resupination of the ovule of the former. That of Ranunculus, if inserted higher, would become hori- zontal ; and if the insertion were transferred to the very summit of the cell, it would be suspended and the rhaphe averse, as in Fig. 581. Upon this conception, Euphorbia and its allies has normally suspended ovules, the rhaphe being next the placental axis, and Buxus and its allies, resupinately suspended ovules, the rhaphe averse. The propriety of regarding the ad- verse rhaphe as the normal condition is confirmed by the fact that the only instance we know of solitary erect ovules from the base of the cell having the rhaphe averse is that of Rhamnus and its allies ; and here it was shown by Bennett (in PI. Javan. Rar. 131), and confirmed by the analyses of Sprague (Gray, Gen. III. ii. 168, plates 163-160), that the rhaphe of the young ovules is ventral, so that the dorsal position, when it occurs, is the result of torsion. J. G. Agardh (in his Theor. Syst. PI. 178, &c.) maintains the contrary, but is not sustained by later observers. Accordingly, even if we adopted Agardh's estimate of the botanical value of the characters here considered, we should prefer to express these differences in the phraseology above indicated, and not to adopt his terms, FIG. 597. Same as 595 more magnified ; the outer coat (a), the inner (6), nucleus (c), and the bundle of spiral ducts (d) in the rhaphe (running from placenta to chalaza) indicated. THE FLOWER. 530. Origin and Nature of the Ovule. It has been already stated in general terms that ovules arc peculiar outgrowths or productions, generally of the margins of carpellarv leaves i .")!.")): thai they are composed of parenchymatous cellular sub- stance, at least as to the nucleus, of which the simplest ovule wholly consists (/>2G) ; that the coats originate subsequcntlv to the nucleus : and that the outer coat is of later origin than the inner one. (•">!*.) The inamiliform protuberance of whicli the forming ovule at first consists originates in one or more cells of a layer directly beneath the epidermis.1 531. The morphological nature of the ovule has been much discussed. The commonly prevalent view was that the ovule is homologous with a leaf-bud, and that its nature is in some degree illustrated by such buds as those which develop on the margins of the leaves of Bryophyllum, as shown in Fig. 322. But such buds, and the bulblets or fleshy. buds which appear on the face of certain leaves, follow the universal order of budding growth, that is, are centripetal in development, the outermost parts being the earlier and the inmost the later formed. The ovule, on the contrary, is basipetal or centrifugal in develop- ment, the nucleus being first and the outer coat last formed; therefore the coats are not homologous with sheathing leaves, nor the nucleus with a vegetative axis. The older theory has accordingly given way to the present one, in which the ovule answers to the lobe of a leaf peculiarly transformed, or to an outgrowth of a leaf, whether from its edges or surface. The u/mtropous, epitropous, and fieterotropous (the first two new, the last employed in ;i new sense), the more so since the application is confused with hypo- thetical considerations and tin- necessity of bringing the ovules ideally back to ascending or horizontal positions. It may be stated, brielly, that //- ti-ii/iiins, iii Agardh's terminology, applies to the normal position of collat- eral ovules, with rhaphes back to back, in opposite directions on the two halves of the placenta; Apotropous, to an erect or ascending ovule witli its rhaplu' next the plaeental axis, ami a hanging one has its rhaphe averse from it : /-'/iiti-0/ions, when an erect or amending ovule has its rhaphe averse, and a hanging one has it adverse. 1 Hofmeister'a statement that the simple ovule of Orchis originates in the division of a single epidermal cell (and is therefore a Iririinmi ) is con- troverted by Strashnrger and by Warming. The latter adds the remark, that even if it were so in cases of extreme simplicity, this would not invali- date the proposition that the ovule is to be regarded as the homologue of the lobe of a leaf. Such a lobe is not rarely reduced to a single bristle. For the whole subject of the origin, development, morphology, and theory of the ovule, see Warming's very elaborate and perspicuous memoir, I )e 1'Ovule; also the papers of (Vlakowsky, Van Tieghem, &c., referred to in notes to paragraphs 500, 501. OVULES. 283 great advantage of this view is that it serves to homologize the fructification of Flowering Plants with that of the higher Flower- less Plants, or the Ferns, the sporangia or analogues of the ovule being outgrowths of the leaf.1 532. Origination of the Embryo. The whole process of fer- tilization and the resulting produc- tion of the embryo, also the history of the subject, belongs to the suc- ceeding volume, involving as they do questions of minute anatomy and of physiolog}'. But a general idea may here be given of the way in which the embiyo originates. The tube which a grain of pollen sends forth into the stigma (574, 575) penetrates the style through loose conducting tissue charged with nourishing liquid, reaches the cavity of the ovary, enters the orifice of an ovule to reach the apex of the nu- cleus, although the latter sometimes projects to meet the pollen-tube. Meanwhile a cavity (the embryo- sac, which is formed by the great enlargement of a single cell of the tissue, or of two or more cells the product of a mother cell) forms in the nucleus, the upper part of it commonly reaching nearly or quite to the apex of the nucleus, which the pollen-tube impinges on or sometimes penetrates. A particular portion of the protoplasm contained in the embiyo-sac forms a globule, and this at the time 1 The advocates of this view naturally maintain that ovules and placentae always belong to leaves, and never truly to a cauline axis ; that in the pro- central placentation of Primulaceae, the actual ovuliferous surface is an out- growth of the bases of the carpellary leaves coalescent with each other and adnate to a prolongation of the torus ; also that in those Gymnosperms which have no earpophyll, such as Yew, the whole nascent carpellary leaf, or rather the papilla which would otherwise develop as such, is directly developed into ovule. This, being solitary and the last production of the axis, necessarily appears to terminate it. (500, 501, notes.) FIG. 598. Diagram representing a magnified pistil of Buckwheat, with longitudinal section through the axis of the ovary and orthotropous ovule ; some pollen on the stigmns. one grain distinctly showing its tube, which has penetrated the style, reappeared in the cavity of the ovary, entered the mouth of the solitary ovule (o), and reached the embryo-sac (s) near the embryonal vesicle (v). 284 THE FLOWER. 604 605 607 of fertilization is found at the apex of the sac, at or adjacent to the part reached by the pollen-tube. Not rarely it adheres to the eoo eoi 602 cos wall of the sac exactly opposite the termination of the pollen-tul ie. This is called the embryonal vesicle. To it the con- tents of the pollen-tube arc in some manner tran — ferred. Upon which it takes a more definite shape, acquires a -wall of cellulose, and so becomes a vegetable cell. This divides into two, the lower again into two, and so on, forming a chain (the sitspensor or pro- I'nihryo). The terminal cell of this divides again and again in three directions, producing a mass of cells which shapes itself into the embryo, the initial plant of a new genera- tion. Ordinarily the sus- pensor soon disappears. It is attached to the ra- dicular end of the em- bryo, which consequently always points to the foramen or microp3'le of the seed. The process in Gymnosperms is more complex, and has to be separately described. 533. Polyembryony, the production of two or more embryos in one seed, is not uncommon in Gymnosperms (there being a kind of provision for it), and is of occasional but abnormal occurrence in Angiosperms. in the seed of Mistletoe, Santahnn, &c. In these it results from the production and fertilization of more than one embryonal vesicle. Strasburger has recently ascer- tained that the commoner polyembryony in the seeds of Onions, Oranges, Funkia. &c., results from the production of adventive embryos, which originate in the nucleus outside of the embryo- sac and wholly independent of fertilization.1 Two kinds of 1 Strnslmrger, TYlier Polyembryonie, in Xritschr. Xntunvis. Jena, xii. 1878 (see Anicr. Jour. Sri. April, 1870). It was found that when, by exclu- sion of pollen, the formation of a normal embryo was prevented, no adventive FIG. 599. Diagram of the suspensor and Incipient embryo af its extremity. 600. The same, with the embryo a litlle more developed. G01. Tlie same, more developed still, tlie cotyledons faintly indicated at tlie lower end. 602. Same, with the incipient cotyledons more manifest. i;n:;. The embryo nearly completed. FIG f.04-fiOr>. Formiiiiremliryo from a half-grown seed of Buckwheat, in three stages. 607. Same, with the cotyledons fully developed. THE FRUM. 285 anomalous reproduction are therefore now known, wkicii are intermediate between sexual and non-sexual, between budding and fruiting propagation, viz.,- Apogamy, which is budding growth or prolification in place of that which should subserve sexual reproduction. This was dis- covered in Ferns by Prof. Farlow, while a pupil of De Bary, by whom our knowledge of the process has recently been extended, and this name imposed.1 The production of bulblets in place of seed or embryo answers to this in Flowering plants. Parthenogeny •, the counterpart analogue of apogani}-, is the non-sexual origination of an embryo extraneous to the embryonal vesicle or even the embryo-sac. However abnormal, its occur- rence is probably not so rare as has been supposed. CHAPTER VII. THE FRUIT. SECTION I. ITS STRUCTURE, TRANSFORMATIONS, AND DEHISCENCE. 534. The Fruit consists of the matured pistil or gyncecium (as the case may be) , including also whatsoever may be joined to it. It is a somewhat loose and multifarious term, applicable alike to a matured ovaiy, to a cluster of such ovaries, at least when somewhat coherent, to a ripened ovary with catyx and other floral parts adnate to it, and even to a ripened inflores- cence when the parts are consolidated or compacted. Fruits, accordingly, are of various degrees of simplicity or complexity, and should be first studied in the simpler forms, namely, those which have resulted from^a single pistil. Such a fruit consists of Pericarp with whatever may be contained in it and incorpo- rated with it. embryo appeared in those seeds which habitually produce them. To this Cffilebogyne offers an exception. The female of this dioecious plant habit- ually matures fertile seeds, with a well-formed embryo, in Europe when there are no male plants in the country. Strasburger ascertained that the embr3ro thus formed is adventive, the embryonal vesicle perishing. Parthenogenesis, of which Caelebogyne was the most unequivocal case, is thus confirmed, and is shown to occur in most polyembryony ; but it is at the same time explained to be a kind of prolification. 1 See Farlow, in Proc. Am. Acad. Lx.68; De Bary ,Bot.Zeit.xxxvi. 465-487. 286 THE FRUIT. 535. The Pericarp, or Seed-vessel, is the ripened ovary. It should, therefore, accord in structure with the ovary from which it is derived. Yet alterations sometimes take place duriim fruc- tilication, either by the abortion or obliteration of parts, or by accessoiy growth. 536. Internal Alterations. Thus, the ovary of the Oak con- sists of three cells, with a pair of ovules in each; but the fruit has a single cell, filled with a solitary seed, only one ovule being mat in fd, while two cells and five ovules are suppressed, the remains of which may be detected in the acorn. The ovary of the Chestnut has six or seven cells, and a pair of suspended ovules in each ; but only one of the dozen or fourteen ovules ever develops into a seed, except as a rare monstrosity. The three-celled ovary of the Horsechestnut and Buckeye is similar in structure (Fig. 608-611), and seldom ripens more than one or two seeds ; but the abortive seeds and cells are obvious in the ripe fruit. The ovary of the Birch and of the Elm is two-celled, with a single ovule in each cell: the fruit is one-celled, with a solitary seed; one of the ovules being uniformly abortive, while the other in enlarging thrusts the dissepiment to one side, and obliterates the empty cell. Similar suppressions in the fruit of parts actually extant in the ovary are not uncommon. 537. On the other hand, then- may be more cells in the fruit than there are primarily in the ovary. Thus the fruit of Datura is dicarpellary and normally two-celled, with a large placenta projecting from the axis far into the cells. But each cell be- comes liilncellate, that is, divided into two, by a false partition growing out from the back of each carpel and cohering with the middle of the adjacent placenta. So the 5-carpellary and nor- mally live-celled ovary of common Flax early becomes spuriously ten-celled (morphologically speaking, not 10-locular. but 10- locellale), by a false partition extending from the back of each FKi. fiKS. Longitudinal s.vtion of the ovary of a ISiirkcvi- (.F.M-nliis I'avia), sl]<>\\ in^ the pairs Of OVUles in two of Ilir rrlls. i;o!l. Transverse section of the s:unr displaying all three cells and six ovules. 610. Same of lialf-^rown fruit, with single fertile s.-,-d, abor- tive ovules and obliterating cells. 611. Dehiscent one-seeded fruit, diminished in size. ITS STRUCTURE AND TRANSFORMATIONS. 287 carpel across its cell (Fig. 539-541) ; and the solitary carpel is similarly divided lengthwise in many species of Astragalus. as in Fig. 534. Transverse divisions or constrictions across o a maturing ovaiy (such as is seen in Fig. 620) are not uncom- mon, especially in legumes and other pods, and are of little mor- phological significance. 538. External Accessions may here be referred to. The wing of the pericarp in Maple, Ash, and the like (Fig. 625-G27) , are familiar instances of this ; and of the same nature are the im- bricated scales which cover some Palm-fruits ; the prickles on the pod of Datura, Ricinus, &c., and the hooked or barbed prickles of many small pericarps (as in various Borraginaceae) , which thus become burs and are disseminated by adhering to the hairy coat of cattle. All these are of the nature of superficial outgrowths, and these especially affect the pericarp or parts connected with it. 539. Persistence of Connected Organs. An adnate calyx (331 ) , being consolidated with the ovaiy, necessarily makes a constit- uent part of the fruit, in the pome (575) doubtless a very large part. The limb or lobes of such adnate organ may persist, as the tips of the sepals on an apple or quince, and may be turned to useful account, as is the pappus of Composite for dissemina- tion. Or, in small pericarps, the style may persist as part of the fruit, and subserve the same ends, either by becoming feathery for aerial dissemination, as in Clematis and in one section of Geum, or by becoming hooked at the tip for adhesion to fleece, &c., as in other species of the latter genus. Or adjacent parts which are not actually incorporated with the pericarp may play similar parts in the economy, as the hooks on the calyx-tube of the dry calyx of Agrimonia, which at maturity is detached with the included fruit, the fleshy fructiferous calyx of Gaultheria (Fig. 651) and of Mulberry (Fig. 654) ; and the pulpy fructiferous re- ceptacle of the strawberry (Fig. 653) : the ultimate utilities in both classes of instances being similar, viz., wide dispersion of the seed by animals, whether by external carriage, or by being devoured and the voided seeds of fleshy fruits thus disseminated. 540. Transformations in Consistence. In the change from ovaiy to mature pericarp, various kinds of transformations may take place. In some the wall of the ovary remains thin and becomes in fruit foliaceous or leaf-like, as in a pea-pod, the carpels of Columbine, and Marsh Marigold (Caltha), or the pod of Colutea or Bladder Senna. In others it thickens and becomes at maturit}' either dry throughout, as in nuts and capsules ; or fleshy or pulpy throughout, as in berries ; or hard-rinded with- THE FRUIT. out but soft within, as in a popo ; or fleshy or berry-like without, but indurated \vitliin, as in all stone-fruits, such as the cherry and i)each. 541. When the walls of a pericarp consist of two laj'ers of dis- similar texture (as in a peach) the outer layer is called EXOCARP, the inner ENDOCARP, these terms meaning exterior and interior parts of a fruit. When the external layer is a comparatively thin stratum or film, it is sometimes termed the EPICARP. When it is fleshy orpulpy it is named SARCOCARP. When the endocarp within a sarcocarp is hard and bony or crustaceous. forming a shell or stone, this is termed a PUTAMEN. When three concentric layers a re distinguishable in a pericarp, the middle one is called Mr>o<- \i;i-. 5-12. Fruits may be divided into two kinds, in reference to their discharging or retaining the contained seeds. They are dehiscent when they open regularly to this end; indehiscent when they remain closed. There is a somewhat intermediate condi- tion, when they rupture or burst irregularly, as in Datura Metel, &c. Dry pericarps with single seeds are commonly indehiscent; those with several or man}' seeds mostly dehiscent. Seeds pro- vided with a wing or coma or any analogous help to dispersion are always in iudehiscent pericarps. Permanently fleshy peri- carps are indehiscent, stone-fruits as well as 1 terries. But in some stone-fruits (/. e., with indurated endocarp and fleshy exocarp), such as those of Almond (Fig. 040) and Hickory, the barely fleshy exocarp or sarcocarp dries or hardens, instead of softening, as maturity is approached, and at length separates from the putamen by dehiscence. 543. Doliiscenco, the opening of a pericarp for the discharge of the contained seeds, is r-gular or fn-ci/ular ; or, better, is normal and abnormal. For most of the abnormal or non-typical modes are as determinate and uniform in occurrence as the typi- cal modes. A good English name for dehiscent pericarps in general is that of POD. 544. Kegular or normal dehiscence is that in which a pericarp splits vertically, for its whole or a part of its length, on lines which answer to sutures or junctions, that is, along lines which correspond to the margins or midribs of carpellary leaves, or to the lines and surfaces (or commissures) of coalescence of con- tiguous carpels. The pieces into which a pericarp is thus sun- dered are termed VALVES. 545. The normal dehiscence of a carpel is by its inner, ven- tral, or ovulifcrou.s suture, that is, by the disjunction of the leaf-margins, as in Fig. 618. Its only other line of normal dehiscence is by the opposite or dorsal suture, that is, down DEHISCENCE. 289 the midrib. Legumes usually dehisce by both sutures (as in Fig. 619), therefore into two valves. 546. A dehiscent pericarp formed of two or more carpels is called a CAPSULE. The two leading terms descriptive of capsular dehiscence were based upon the modes of opening of pericarps having as many cells as carpels : they are the septicidal, that is, as the term denotes, cutting through the septa or dissepiments ; and the loculicidal, that is, cutting into the loculi or cells. 547. Septicidal, the dehiscence through the dissepiments, is the disjunction of a pericarp into its constituent carpels, these then usually themselves dehiscing down their ventral suture, as in Fig. 612, illustrated by the diagram, Fig. 613. Good examples are furnished by the Hypericum Family (the pistil illustrated in Fig. 536, 537), where the placentae which compose the axis are carried awa}^ on the edges of the par- titions or introflexed valves ; also by Rhododendron, Kal- mia, and the like, in which the placentae remain combined into a column in the axis (the COLUMELLA or column) , from which the edges of the valves break away. 548. The septicidal dis- junction of the carpels does not of itself open the cells. Such separated carpels when one-seeded not rarely remain closed, as in Mallow, Ver- bena, &c. Or when dehiscent they may open both by the ventral and dorsal sutures ; i. e. , the pericarp may first divide into its constituent carpels, and then each carpel break up into half carpels, as in Euphorbia. 549. Loculicidal, the dehiscence into the loculaments, loculi, or cells of the pericarp (shown in Fig. 614, and the diagram, 615), is that in which each component carpel splits down its FIG. 612. Septicidally dehiscent tricarpellary capsule of Elodes Virginica. 613. Dia- gram of septicidal deliiscence. FIG. 614 Loculicidally dehiscent tricarpellary capsule of an Iris, divided trans- versely at the middle. 615. Diagram of loculicidal dehiscence. 290 THE FRUIT. dorsal suture, as in Iris, Hibiscus, CEnothera, &c. In this, the dissepiments remain intact. If they break away from the centre then they are borne on the middle of the valves, as in the figures above cited. If they remain coherent in the axis but break away from the valves, the result is one form of what is called - 550. Septff nii/nl dehiseence, i. e., a breaking away of tin- valves from the septa or partitions, as shown in Fig. GIG. This represents the loculicidal form of the sept i frugal mode, which is less common than that of the accompanying diagram, Fig. G17. Here the partitions alternate f — X,, /^~ s. with the valves: that is. the dehiseence of the pericarp is I \ i \v^^x^^ i of the septicidal order, as I .^''^ \^ \ I ) near as may be. but the par- \ / \ J titions do not split, wherefore Vs, ^S N^ ^/ the valves break away at the (!1G 617 common junction. To this the term marginicidal has been applied. It occurs in the 2-3- carpellary capsule of Ipomoea (especially in the common Morning Gloiy), in the 5-carpellary capsule of the North American species of Bergia ; likewise in the 2-carpellary pod of Crucifene (Fig. 623), with a difference that the placenta- from which the valves break away are here parietal and the partition is abnormal. 551. The terms septieidal and loculicidal apply equally in plan, though not with etymological correctness, to one-celled capsules with either parietal (-195) or free central (599) placentae. AYhcn the dehiseence is of the septicidal type and the placcntation pari- etal, the (half) placentae are borne on the margin of the valves, as in the Gentian family and the species of Ilypericum with one- celled capsule. When the placenta- are borne on the middle of the valves, as in Violets, the dehiseence is of the loculicidal type. In the case of free central placentae with no trace of partitions, the character of the deliisceiice may usually be deter- mined by the position of the styles or stigmas relative to the valves. .").") 2. Dehiseence may be quite normal although very partial, as when confined to the apex of the capsule of Cerastium and of rrimula, and even to the pores under the radiate stigmas of Poppy. .">:>:',. /my/»/w or , ;i> to primary kinds, is into 1. Nuts, or Achcenocarps, dry and indeliisci-nt ; 2. Pods, or Rerjmacarps, dry, dehistvnt ; 3. Stow -/',-n/is, or Fi/rrnonirps, flesliy without, indurated within, indehiscent; 4. I* rr/fs. or Sarcocarps, floshy throughout, indc'liiso'iit. FIG. f.is. Adelilacent follicle of Marsh-Marigold, Caltha i>ulustris. FIG. C19. Legume of a Sweet Pea, already dehiscent. C20. Loment of a Desmodlum. ITS KINDS. 293 which that order presents. Some of these, in fact, are in- dehiscent and reduced to akenes ; some break up at maturit}r into one-seeded indehiscent articulations or joints, which are dispersed as if they were so many seeds. A legume of the latter kind takes the special name of LOMENT, Lat. Lomentum. (Fig. 620.) In Mimosa (Sensitive-plant, &c.), such articulations de- hisce into two valves. They also fall away from the sutures, or from a persistent marginal border of them, or in some cases the valves thus fall away entire. The persistent frame which remains has been called a REPLUM, an architectural word, here taken in the sense of door-case. 559. A Capsule is the pod, or dehiscent fruit, of any compound pistil. When regularly and com- pletely dehiscent, as already stated (544), the pod splits lengthwise into pieces or voices. The modes of regular dehiscence are illus- trated in Fig. 612- 617. Two modifica- tions of the capsule have received distinc- tive names which are in common use, viz. the Pyxis and the Silique. 560. A Pyxis or Pyxidium is a dry fruit which opens by a circular line, cutting off the upper part as a lid ; i. e., the dehiscence is circumscissile. (553, Fig. 621 .) In the Purslane, Pimpernel, Henbane, and Plantain, the pyxis is a capsule ; in Amaranths (Fig. 637) it is a utricle; in Jeffersonia (Fig. 622) it is a modi- fication of the follicle, being of one carpel which dehisces transversely, and not all round, so that the lid remains attached. 561. A Silique is a narrow two-valved capsule, with two pari- etal placentae, from which the valves separate in dehiscence ; as in plants of the Cruciferous or Mustard family (Fig. 623), to the fruit of which this term is restricted. Usually, a false partition is stretched across between the two placentae, render- FIG. 621. Pyxis of Purslane, Portulaca oleracea, the top separating entirely and falling away. FIG. 622. Pyxis-like follicular fruit of Jeffersonia diphylla; the lid remaining attached dorsally. FIG. 623. Silique of Cardamine. in dehiscence. 624. Silicle of Capsellaor Shepherd's- Purse, lateral view, and an oblique view of the same with one valve removed. 623 294 THE FRUIT. ing the pod two-celled in an anomalous manner. A SILICLE (Xillcula, diminutive of x!ln/n">."•) an- also akcncs. Tho name is extended to all one-celled seed-like fruits resulting from a FIG. 625. Samara or key of White Asli. Fraxinus Americana. C.L'ii. That of White Elm. Ulinns Amcrir:m:i C,-J7. h.uil'lr s:iin:ir:i of KIM 1 Majili-. Acvr rulinmi. !•'!<;. I;L'.S. Acheiiiuni of a common Buttercup. 629. Vertical sec tioii, showing the seed within. ITS KINDS. 295 compound ovary, and even when invested with an adnate tube. Of the latter is the fruit of Composite. (Fig. 630 Here the tube of the calyx is incorpo- rated with the surface of the ovary ; and its limb or border, obsolete in some cases (Fig. 630), in others appears as a crown or cup (Fig. 631), or set of teeth or of scales (Fig. 632, 633), or as a tuft of bristles or hairs (Fig. 634, calyx- -635.) 635), &c., called the PAPPUS. In the Lettuce and Dandelion (Fig. 635), the achenium is rostrate, or beaked, i. e. its summit is extended into a slender beak. An akene with adnate ealyx has been termed a CYPSELA. 565. The Utricle is the same as the akene, only with a thin and bladdery loose pericarp, like that of Goosefoot. (Fig. 636.) This thin coat sometimes bursts irregu- larly, discharging the seed. In the true Amaranths, the utricle opens by a circular line, and the upper part falls as a lid, converting the fruit into a small pyxis (560), — a transition form. (Fig. 637.) 566. A Caryopsis or Grain differs from the utricle or akene in having the seed completely filling the cell, and its thin coat firmly consolidated throughout with the very thin pericarp ; as in wheat, Indian corn, and all other cereal grains. Of all fruits this is the kind most likely to be mistaken for a seed. 567. A Nut is a hard, one-celled and one-seeded, indehiscent fruit, like an achenium, but larger, and usually produced from an ovary of two or more cells with one or more ovules in each, all but a single ovule ' and cell having disappeared during its growth (536) ; as in the liazel, Beech, Oak (Fig. 638), Chest- FIG. 630. Achenium of Mayweed (no pappus). 631. That of Cichory (its pappus a shallow cup). 632. Of Sunflower (pappus of two deciduous scales'). 633. Of Sneezeweed (Helenium), with its pappus of five scales. 634. Of Sow-Thistle, with its pappus of delicate downy hairs. 635. Of the Dandelion, tapering below the pappus into a long beak. FIG. 636. Utricle of Chenopodium album, or common Goosefoot. 637. Utricle of an Amaranth, by transverse dehiscence becoming a pyxis. 296 THE FRUIT. nut, and the like. The nut is often enclosed or surrounded by a kind of involucre, termed a <'tij>nle; such as the cup at the base of the acorn, the bur of the ehe.stimt. and the leaf- like covering of the hazel-nut. The name (,'///„* (sometimes (ilmul in Knglish) is technically applied to such nuts, this being their classical Latin name. 5('>71. The pair of achenium-likc or often samara-like carpels, Nut and akene. bet ween which there is no fixed distinction, will cover this ground. The fruit of Cyperacese, for instance , is truly an achcnium, if this name is ever to be used (and it now commonly is) for any other than a monocarpellary fruit. It is often termed a nut, sometimes a nutlet, and by a late writer, Bcrekler. a caryopsis. '-' ('. ITS KINDS. 297 united by their inner face but separating entire at maturity, which constitute the fruit of Umbelliferae, takes the name of CREMOCAKP (Lat. Cremocarpium) ; arid the halves are called MERICAKPS. These names it may sometimes be convenient to use ; yet it is not advisable to have special names for the fruits of particular families; and mericarp is here synonymous with carpel. For dry fruits in general (or such as become dry) which are composed of two or more carpels, and which at matu- rity split up or otherwise separate into two or more closed one- seeded portions, an appropriate recent name is that of SCHIZOCARP. The component carpels of such a fruit were long ago named Car- cerules (carceruli, little prisons) b}r Mirbel. 572. Fleshy Fruits, which from their texture are naturally indehiscent, may be either fleshy throughout, or with a firm rind or shell, or fleshy externally and hard or stony internally. Of the latter, the type is 573. The Drupe or Stone Fruit proper (Fig. 639), that of the cherry, plum, and peach. True drupes are of a single carpel,, one-celled and one-seeded (or at most two-seeded) , in /. \ "V the ripening of which the . outer portion of the pericarp becomes fleshy or pulpy, and the inner stony or crustace- ous, i. e. divides into sarco- carp and putamen. (541.) But the name is extended to pericarps of similar texture i-esulting from a compound pistil, either of a single cell, as in Celtis, and (by abortion) in the olive, or of two or several cells, as in Cornus, Rhnmnus, &c. The several pericarps of the aggregate blackberry and raspberry are diminu- tive drupes or DRUPELETS. 574. Small drupes are often confounded with berries, and the stone or stones taken for seeds. Especially is it so in drupes or drupaceous fruits of more than one cell, ripening into separate or separable hard endocarps or stones, each filled b}r a seed.1 Bearberries (Arctostaphylos) and Huckleberries (Gaylussacia) are good illustrations of this. The seed-like endocarps of this 1 The term Acinus, the original name of such a berry as a grape, hns been used in descriptive botany for a small drupe or drupelet, and the ripened carpels of TCubus have been termed acini or acines, but without discriminating them from berries. FIG. 639. Vertir.nl section of a peach. 640. An almond ; in which the exocarp, the portion of the pericarp that represents the pulp of the peach, remains juiceless, and at length separates by dehiscence from the endocarp, or shell 298 THE FRUIT. sort are PYREN^ ; and the fruits are dipyrenous, tripyrenous, tetrapyrenous , &c., according as they contain two. three, or four pyrenae. When the sarcocarp is thin and dries up at maturity, these p}Tenae pass by gradations into nuculae (5G'J) or nutlets: hence pyrenae are not uncommonly in English descriptions called nutlets or nucules. 575. The Pome (Fig. 641, 642) is the name of the apple, pear. and quince. These are fleshy fruits, composed of two to several carpels (rarely by abortion only one)- <>f parchment-like or (in Hawthorns) bony texture. enclosed in flesh which morphologically belongs to adnate calyx and receptacle ; as may lie ap- prehended by comparing a rose-hip (Fig. 407, in flower) with an apple or a pear. Of the quince, the whole flesh is calyx or hypanthium (39.")) ; in the apple and pear, the inner or core-portion of the llesh is of the nature of disk, investing the carpels. In the fruit of Hawthorns, the carpels become bony pyren;e (574), and so the fruit is drupaceous, is indeed nothing more than a syncarpous drupe. In Eriobotrya, or Cumquat, the carpels becoming veiy thin and membranaceous, the pomaceous fruit is in fact a kind of berry. 576. The I'epo, or Gourd-fruit (Fig. 643), of which the gourd and squash are the type, and the melon and cucumber equally familiar illustrations, is the char- acteristic fruit of Cucurbitacese, llesh}" internally and with a hard or firm rind, 11 or part of which is referable X) the adnate calyx completel1 incorporate with the ovary. This is either one-celled with three broad and re volute parietal placenta1, or these pla- centae, borne on thin dissepiments, meet in the axis, enlarge, and spread, unite with their fellows on each side, and an- rellccted to the walls of the pericarp, next which they bear their ovules. As the fruit enlarges, the seed-bearing placenta- usually cohere with the walls, and the partitions are obliterated, giving the FI(;. 641. Pome or apple in transverse section. C.rj. (.miner in vertical section : the inner flesh answering to disk in tin- apple and pear is liere wanting. FIG. 643. Seel ion of the ovary of the Gourd. 644. Diagram of one of its constituent carpels. I'll ITS KINDS. 299 appearance of a peculiar abnormal placentation, which the study of the ovary readily explains. In the watermelon the edible pulp all belongs to the greatly developed placentae. Fruits of this family in which the rind also is soft at maturity are true berries. 577. The Hesperidium (orange, lemon, and lime) is the fleshy fruit of a free many-celled ovary with a leathery rind, and is a mere variety of the berry. The name is applied only to fruits of the Orange tribe. 578. The Berry (Lat. Bacca) comprises all simple fruits in which the pericarp is fleshy throughout. The grape, gooseberry, currant, cranberry (Fig. 645), banana, and tomato are familiar examples. The first and last consist of an ovary free from the calyx ; in the others, calyx and ovary are combined by adnation. 579. Aggregate Fruits are those in which a cluster of carpels, all belonging to one flower, are crowded on the receptacle into one mass, as in the raspberry and black- berry taken as a whole. (Fig. 646.) They may be aggregates of any kind of simple fruits. But when dry and not coherent, the mass would simply and properly be described as a head or spike of carpels, more commonly of akenes, as in Ranunculus, Ane- mone, &c. Yet when numerous carpels thus compacted become fleshy, and sometimes more or less coherent, the aggregate ma}' need to be taken into account. The best name for it is that of SYNCARPIUM, or in English form SYNCARP. But the term has been applied to multiple fruits as well.1 In Ilydrastis, the numerous carpels imbricated on the upper part of the torus arc baccate, that is, become berries ; in a raspberry, 64? the seemingly baccate grains are drupaceous (being drupelets, 573), 1 The sijttcarp which is a gyncecium might be designated a sim/>/<> s//n- carpium : that which is an inflorescence, a complex syncarpium, which may be biflorous, pauciflorous, or multiflorous. FIG. 645. The larger Cranberry, Vaccinium (Oxycoccus) macrocarpon ; the berry transversely divided. FIG. 646. Vertim! section of half of a blackberry) of RubusvUlosus), enlarged; and, 647, of one of its drupelets more magnified. f 300 THE FRUIT. and. slightly cohering together (though vrithout organic union), they tail as OIK- body from UK- conical dry torus at maturity. It is the same in blackberries or brambli --berries 0), such carpels, imbricated over one another, cohere inure or le-- at all contiguous pails, and become drupaceous ; never- theless, at maturity each opens dorsally. allowing the seeds to fall out : in age it dries and hardens, and also separates from its connec- tions, and so be- comes a follicle, but with the remark- able peculiarity of dorsal instead of ventral dehiscence. (Fig. C")0.) In Li- riodendron. a tree of the same family, such carpels are dry and indehiscent throughout ; and they largely consist of long and flat styles, imbricated in a cone, but separating from each other and from the slender torus at maturity, when each becomes a samara. 580. Accessory or Anthocarpons Fruits are those of which some conspicuous portion of the fructification neither belongs to the pistil nor is organically unite. 1 with it. except by a common insertion. The part thus imitating a fruit, while it is really no part of the pericarp, is sometimes called a Pseudocarp, or an .\nthocarp or Anlhocarpium. This condition may occur either in simple, in aggregate, or in multiple fruits. 1 Tin' aT'.nvMtc fruit like that of Kubiis (named by some Conocar/iiiiin, by others an . /'/< rio, Kri/tJtroxtoniuin, &c.) was termed by Dumortier a Drupe- Inn,. A similar aggregation of baccate carpels lie termed a lla,;;lnm ; of follicles, a FolKcetum, &<•. All such names may look well in a system; but they are both superfluous and unmanageable in phytography. FIG. G4S. A"!_'r«"::it,' fruit of Unil>rpll!i-trn>. Magnolia tTinbrrll:i. n-.luoed in size; a seed from a lower .l.-Viscrn' .-arpd linngs on a tlnvad (•diiMstini: of a tnft of extensile spiral ducts iinr.avrllc.1. C.J'.i. Same in ]onirituilin:il M-i'tii.n. C.r.n. one ..f tin- carpels detached, .-t full maturity, dried up, dorsally dehiscent, exposing the pair of seeds of the natural size. ITS KINDS. 301 581. Gaultheria procnmbens, the aromatic Wintergreen (Fig. 651, 652), affords a good example of the first. Its seeming berry (the checkerberry) , with summit crowned by the tips of the calyx-lobes, well imitates the true berry of a Vaccinium, such as that of Fig. G45. But it comes from a flower with thin calyx, underneath and free from the ovary. Its fruit is really a capsule : in the process of fructi- fication, the calyx enlarges, becomes succulent, completely encloses the capsule or true fruit, yet without adhering to it, and in ripening counterfeits a red berry. So in Shepherdia, or Buffalo Berry, the seeming sarco- carp of a drupe is really a free calyx, accrescent and succulent, enclosing an akene. So, also, the apparent acheniurn or nut of Mirabilis, or Four-o'clock, and of its allies, is the thickened and indurated base of the tube of a free calyx, which contracts at the apex and encloses the true pericarp (a utricle or thin akene) , but does not cohere with it. 582. Likewise the torus, although not con- spicuous, may be said to be an accessory part of the aggregate fruit of the Blackberry or Bramble (579) : it becomes the solely con- spicuous and the sole edible part of a straw- berry (389, Fig. 406, 653), the akenes or true fruits dispersed over the surface being apparently insignificant. Equally in man}' multiple fruits the conspicuous flesh belongs to receptacle (either torus or rhachis), to calyx, or even in part to bracts, or to all these parts combined, as in a pine-apple. 583. Multiple or Collective Fruits l are those which result from the ao-o-regation of several flowers into one mass. The simplest OCT O of these are those of the Partridge-Berry (Mitchella, Fig. 467), 1 Collective is the preferable name. The term multiple was applied by DeCamlolle to what are here (following Lintlley) called a^jrcijtitf fruits ; and the (tyr/reyate fruits of DeCandolle are here called multi/>/< or collect ice. Moreover, the distinction between accessory or anthocarpous and collective or multiple fruits was not recognized by Lindley, who combined the two in his original " Introduction to Botany." In this work four classes are given : 1. Fruit simple, APOCARPI ; 2. Fruit aggregate, AGGREGATI ; FIG. 651. Forming capsule of Gaultheria procumbens. with enlarging calyx partly covering it. 652. Same, more advanced, and in longitudinal section . FIG. 653. Vertical section of half a strawberry. Compare with Fig. 406. 302 THE FRUIT. and of certain species of Honeysuckle, formed of the ovaries of two blossoms united into one fleshy fruit. The more usual sorts are such as the pine-apple, mulberry, and the fig. These arc. in lad. dense forms of inflorescence, with the fruits or Moral envelopes matted together or coherent with cadi other: and all or some of the parts succulent. The grains of tin- mulberry (Fig. (',:>[-('>:>('>) are not the ovaries of a singJe flower, like those 058 657 655 656 of the blackberry, which it superficially resembles: they belong to as many separate Mowers : and the pulp pertains to the calyx, not to the pericarp, which is an akene. So that this, like most multiple fruits, is anthocarpous as well as multiple. Similarly, the mostly indefinite fructiferous masses of Strawberry Elite may resemble strawberries: but the pulpy part is the calyx of many Mowers, not the succulent receptacle of one. In the pine-apple, the Mowers arc spicate or capitate on a simple axis, which grows on beyond them into leafy stem; this when rooted as a cutting :;. Fruit compound (ovaria compound), SYNCARPI ; 4. Collective fruits, . loiter, in his "Elements of Rot.-iuy." I.imllev reduced tlie classes to two: 1. Siiiif/i' /•>///'/*. tlmsc prne.cedinir from a single flower; 2. Multifile i/lfi, those formed out of several flowers. FIC. r,.->J. A mulberry. young. 055. One of the tlcsliy grains :il (lowering time, show- in •,' it lo !>'• n i'i.". A yoiini; tig. f.r>S. Longitudinal section of the s;unc later, but in flowering time. !>.">!>. A small slice, magnified, showing some of the flowers. ITS KINDS. 303 bears another pine-apple, and so on : the constituent flowers have through immemorial propagation in this way become sterile and seedless, and all its parts, along with the bracts and the axis of the stem, blend in ripening into one fleshy and juic}" mass. Few fruits of this class have ever been technically named, at least with names which have come into use. But the two following deserve special appellations, although only the latter is familiar either in ordinary language or in descriptive botan}'. 584. The Syconitim or Hypaiithodium, the Fig fruit. (Fig. 657- 65(J.) This results from a multitude of flowers concealed in a hollow flower-stalk, if it ma}* be so called, which becomes pulpy and edible when ripe ; and thus the fruit seems to grow directly from the axil of a leaf, without being preceded by a blossom. The minute flowers within, or some of them, ripen their ovaries into veiy small akenes, which are commonly taken for seeds. The fig is to the mulberry what a rose-hip is to a strawberry. (389, Fig. 406, 407.) It is further explained by a comparison with a near relative of the Fig-tree, Dorstenia, in which similar flowers cover the upper surface of a flat peltate disk. This disk or plate sometimes becomes saucer- shaped b}* an elevation or incurvation of the margin. A greater degree of this would render it cup-shaped, or even pitcher-shaped ; from which it is a short step to the contraction of the mouth down to the smaU orifice which is found in the fig. 585. The Strobile or Cone (Fig. 660) is a scaly multiple fruit, resulting from the ripening of certain sorts of catkin. The name is applied to the fruit of the Hop, where the large and thin scales are bracts ; but it more especially belongs to the Pine or Fir cone, the peculiar fruit of Coniferae (507), in which naked seeds are borne on the upper face of each fructiferous scale (Fig. 661), or some- times in their axils. Such a cone when spherical, and of thickened scales with narrow base, as that of Cypresses, has been termed a a'° GALBULUS, an unnecessaiy name. The galbulus of Juniper is a FIG. 660. Strobile or Cone of a Pitch Pine, Pinus rigida. 661. Inside view of one of the scales, showing one of the winged seeds, and the place from which the other, 662, has been detached. 304 THE FRUIT. remarkable transformation into a seeming berry ; the few scales cohering with each other as they grow and becoming fleshy at maturity, completely enclosing a few bony-coated seeds. :>si;. A Synopsis of the kinds of Fruit, as characterized in this chapter, is appended. The analysis extends only to simple fruits. For there are no commonly used special names of kinds of Aggregate (579), Accessory (580), or Multiple (583) fruits, except that of Strobile. SIMPLE FRUITS are Dry and dehiscent, monocarpellary, Opening \>y one (chiefly the ventral) suture, FOLLICLE. Opening by both sutures, LBGDMJB. Or transversely jointed, LOMEXT. Dry and dehiscent, bi-pluri-carpellary, CAPSULE. When its dehiscence is circumscissile, PYXIS. When dehiscent by two valves from two parietal placentae, . . SILIQUE. A short and broad silique, SILICLE. Dry and bi-pluri-carpellary, splitting into one-seeded carpels, . SCHIZOCARP. The dimerojus schizocarp of Umbellifera, CREMOCARP. Kach of its halves or carpels, HEMICARP or MERICARP. The akene-like or nut-like parts into which Schizocarps generally divide, NUCULES or NUTLETS. Dry and indehiscent, one-celled, one-two-seeded, Winged SAMARA. Wingless, and with the Thin pericarp consolidated with the seed, CARYOPSIS Thin pericarp loose and not filled by the seed, UTRICLE. Thick or hard pericarp free from the seed, Small, from a one-celled one-two-ovuled ovary, AKENE or ACHENIUM. Larger, mostly from a two-several-celled and ovuled ovary, . NUT. Nut borne in a eupule or involucre, GLANS. Fleshy and indehisceiit, Heterogeneous in texture, having A stone (put.-iineii) or nutlets within an exterior sarcocarp, . DRUPE. Papery or cartilaginous carpels in an inferior sarcocarp, . . POME. A harder or firm rind or exterior, and soft interior, From an inferior ovary (confined to Grourd Family), . . • PEPO. From a superior ovary (confined to Orange Family), HESPERIDIUM. Homogeneous, fleshy throughout, BERRV. THE SEED. 305 CHAPTER VIII. THE SEED. 5i'.7. THE SEED is the fertilized ovule (515), with embryo formed within it. It consists, like the ovule, of a nucleus or kernel, enclosed by integuments. The seed-coats are those of the ovule, viz. two, or sometimes only one, in certain plants none. Occasionally an accessory coat appears after fertiliza- tion ; and certain appendages may be produced, as outgrowths from some part of its surface or from its base. The nucleus or kernel is composed either of the embryo alone, or of a nutritive deposit in addition. (19-41.) All the parts of a seed are in- dicated in Fig. 663. o.S.S. The SEED-STALK or PODOSPERM, when there is one, is the faniculus of the ovule (516), and retains this name. So also do the CHALAZA, RHAPHE, and HILUM ; the latter being the scar left by the separation of the seed from its funiculus or directly from the placenta. The foramen of the ovule, now closed, is the MIOROPTLE of the seed. 589. The terms which denote the char- acter of the ovule, such as orthotropous, campylotropous, amphitropous , and anatropous, apply equally to the resulting seed. 590. Seed-Coats. The integuments of the seed answer to the primine and secundine of the ovule. The main seed-coat is the exterior integument of the ovule when there is more than one. Being the most firm coat, and not rarely crnstaceous in texture, it takes the name of TESTA, which is equivalent to seed-shell. It has also been named SPERMODERM (seed-skin), and sometimes Episperm. The latter name (meaning upon the seed) is best applied to the pellicle or outer layer, sometimes a thick one, which the testa of certain seeds forms. The testa is extremely various in form and texture, is either close and conformed to FIG. 663. Vertical magnified section of the (anatropous) seed of the American Lin- den; with the parts indicated, viz. the hilnm (a); testa (6); tegmen (r); albumen ('/); embryo (e). 664. Vertical section of the orthotropous seed of Heliauthemuia Cana- dense, with its funiculus, a. 306 Till-; SEED. the nucleus, or loose and cellular (as in Pyrola-seeds) , or vari- ously appendaged. •V.il. The inner coat, called TEGMKN and sometimes ENDO- PLEUKA, when present is always conformed to the nucleus, and is thin or soft and delicate. .Sometimes it is inconspicuous through cohesion with the nucleus or with the inner surface of the testa. In ovules of one coat it is necessarily wanting. 592. Appendages or outgrowths of the testa generally have reference to dissemination. Two characteristic kinds of such appendages are the wing and the coma. both pertaining only to the seeds of dehis- cent fruits and calculated, by rendering seeds buoyant, to facilitate dispersion by the wind. The wing of a Pine-seed (Fig. 661, 662) is a part of the carpellary scale upon which the two ovules grew. In Trumpet Creeper (Fig. 665), an entire wing surrounds the body of the seed. In the related Catalpa (Fig. 666), it is mainly extended from the two ends, and almost dissolved into a coma, the name given to the tuft of soft hairs like that which forms the down at one end of the seed of Milkweed (Fig. 667), and of Fpilobimn, and at both ends in several Apocynacea?. In the Cotton-plant, very long and soft hairs, admirably adapted for spinning, thickly cover the whole seed- 666 coat. The wing and coma of seeds are functionally identical with the wing and the pappus of the pericarp in the samara and the akenes of Composite (">63, 564), but morphologically quite unlike them. 593. There are other (mainly microscopic) structures on some seed-coats which come usefully into play in arresting farther dispersion al a propitious time or place. In many but not all Polemoniace;e (notably in Collomia), in certain Acanthace;e. such as IJnellia tuberosa (and equally in certain Composite of the Scnccio tribe and in Salvias, &c., among Labiatie. where this structure is transferred to akenes and nutlets), the testa is coated with short hairs, which when wetted burst or otherwise open and discharge along with mucilage one or more very atten- Fic. r.r>r>. Winged seed of Trumpet Creeper, Tecomaradicana <;r,r,. That of Catalpa, becoming cniuosr: tin- l»nly iliviih'cl Irn^tlnvisi- tlm>tii,'h the embryo. FIG. 667. Comose seed «>1 Milkweed, Asclepias Cornuti. ITS COATS AND APPENDAGES. 307 d uated long threads (spiricles) which were coiled within. These, protruding in all directions and in immense numbers, form a limbus of considerable size around the seed, and evidently must serve a useful end in fixing these sma1! and light seeds to the soil in time of rain, or to moist ground, favorable to germination. In cress and flax-seed, the abundant mucilage developed when wetted comes from the gelatination of epidermal cell-walls, and subserves a similar use. 594. While the testa in many seeds is hard and crustaceous or boiry, imitat- ing the pericarp of a nut, in others (such as Paeonia) it becomes berry-like (baccate), and in Magnolia, drupaceous.1 (Fig. 668- 671.) These may also be regarded as adaptations for dissemi- 672 nation, here b}- the agency of birds, attracted ~by bright coloring and edible pulp. 595. The rhaphe of an anatropous seed (shown in Fig. 681, 685) is sometimes so salient as to form a conspicuous appen- dage, as in Sarracenia, Fig. 672. Again it may be wholly 1 See article On the Structure of the Ovule .and Seed-coats of Magnolia, in Jour. Linn. Soc. ii. 100, from which the accompanying figures and Fig. 589-597 are reproduced. FIG. 668. Forming seed (one eighth of an inch long) of Magnolia Umbrella; the rhaphe toward the eye. GG9. Magnified view of the same divided lengthwise through the rhaphe; the outer coat, a. beginning to form a hard inner layer. nf . Within and distinct from this is the inner coat (l>\ immediately enclosing the nucleus, c. The oppo- site side of the testa is thicker on account of the rhaphe, in which d indicates the cord of spiral ducts. FIO-. 670. A nearly full-grown seed, of the natural size 671. Longitudinal section, enlarged, snowing the crustaceous or stony inner stratum of the testa well developed: the parts lettered as in Fig. C>G!>. 672. A transverse section in the same position. 308 THE SEED. inconspicuous, as in the ripe seed of Magnolia, -where it is at length completely merged mid imbedded in the fleshy drupaceous tc^ia, us shown in Fig. 670-07:.'. ;V.H>. Crest-like or other appendages are not uncommon either on the rhaphe or at the hiliiin. These are outgrowths produced (luring the development of the ovule into the seed. In Sangui- naria, such a crest develops from the whole length of the rhaphe C73 D74 (Fig. 673) ; in Dicentra, Corydalis (Fig. G74), &c.. from some part of it, mostly from its base next the hilum. or from the hilum itself, or even from just below it. Such an appendage, especially when attached to the base of the seed, is named a STROPHIOLE. A similar and commonly a wart-shaped appendage in Euphorbia, Riciuus (Fig. G7">). &c., is prodneed by an out- growth of the external orifice of the ovule, the micropyle of the seed. This properly takes the name of CAIHXCLK. But the two terms are not always discriminated. By further develop- ment, cither of these in 33* give rise, in certain seeds, to an acces- sory covering called 597. The Aril or Arillns. This term, rather vaguely employed Ity Linnaeus, was first well defined b3r Gaertner. The true 73. Jviine of S!iiejuiii:iri:i or Hloodroot. with rhaphe crested for its whole length. I'M I Seed of Corydalis aurea, with erest. or strophinlc, attached at or near the hilum. 675. Seed (suspended1) of Riciuus. with its raruncle. PIG. 671!. Seed of White Water-I.il y, Nynijilisea odorata, in its loose and thin arillus. ere ARILLUS, ALBUMEN. 809 the pulp of the berry consists of these fleshy arils, much com- pacted. (Fig. G77, G78.) 598. The laciniate aril of the nutmeg (mace) and, it is said, the bright red and pulp}' aril of Euonymus and Celastrus begin in the manner of a ca- runcle, and are formed (mainly if not wholly) of an outgrowth at or around the micropyle. So that, if an orthotropous seed ever developed an aril of this sort, it would 6-7 ers be seen to begin at the apex of the seed and cover it from above downward. Planchon, who distinguished this from the true aril, gave to it the name of ARILLODE (Arittodiuni) or False Arillus. 599. The Nucleus, or kernel of the seed, consists of the Albu- men, when this substance is present, and the Embryo. 600. The Albumen, as described in the second chapter (25, &c.) , is the name generally employed b}T s}'stematic botanists for a store of nutritive matter in the seed outside of the embryo, whatever its chemical composition. It is not here the name of a chemical substance (albumen or albumin), but of a cellular structure, the cells of which arc loaded commonly with starch- grains (as in the Cerealia), more or less mingled with other matters, or else filled with an encrusting deposit of some equiva- lent substance, as in the cocoanut, coffee-grain, &c. The cells in which this deposit is made belong either to the original tissue of the nucleus, or to a new formation within the embryo-sac, mostly to the latter. (503.) 601. Albumen ma}- be said to belong to all seeds in the grow- ing stage. In what are called albuminous seeds it persists and forms either almost the whole kernel, the embryo remaining minute (as in Fig. '23, 54, 680), or forms a large portion of it (Fig. 13, 17, 19, 21, 48, 663, 664), or, by the growth of the embryo displacing it, it may in the ripe seed be reduced to a thin stratum or mere lining to the contiguous seed-coat ; or it ina}- disappear altogether, as in the seeds of Maple, Almond, Squash, Pea, and the like, which are therefore said to be exalbuminoits. The difference between albuminous and exalbuminous seeds is that the maternal nutritive deposit is transferred to the embryo in FIG. 677. Section of pericarp and placenta of Podophyllum peltatnm; the pulp of the latter mainly of the nature of arillus. investing the seeds. 678. The arillus of one seed detached and enlarged, divided lengthwise, showing the seed within. 310 THE SEED. the former during germination, in the latter during the growth of the seed. 602. The albumen was named Perisperm by Jussieu, and Endosperm by Richard (2/J, note) ; but neither name has in sj'stematic botany displaced the earlier one of Grew and Ga.-rt- ner. But both names have recently been brought into use to distinguish between two kinds of albumen, that formed within the embryo-sac, which is specifically termed KM>OMM.I;M. and that formed without, which takes the name of PEIJISPKKM. This use comports with the etymology of the two words, the former refer- ring to a comparatively internal and the latter to an external portion of the seed or kernel. di):}. In most seeds the albumen is endosperm: in Canna it is all perisperm. In Xympluea and its allies (except Xelum- bium, which has none) mostof it is perisperm ; but a thin and condensed layer of endosperm surrounds the embiyo, where with the per- sistent embryo-sac (or the apex of it) it forms the fleshy sac in which the embryo is enclosed. It is the same in the Pepper Family (Fig. 679) , except that there is a larger quau- <;r:i tit}* of endosperm or inner albumen. 604. When the nucleus of a ripe seed is hollow, as in the cocoanut and mix vomica, the formation of endosperm, which usually begins next the wall of the embryo-sac, has not proceeded so as to fill the cavit}'. The embryo-sac in the cocoanut attains enormous size, and the cavity is filled by the milky fluid. 60"). The texture or consistence of the albumen di tiers greatly. It is I'ltn'tKiceous or mealy when, consisting mainly of 'starch- grains, it may readily be broken down into a powder, as in wheat, buckwheat, &c. ; "////. when saturated with a fixed oil, as in poppy-seed; Jleshy, when more compact, but readily cut with a knife, as in thi! seed of Barberry; mttrifiti/i/inns, when soft and somewhat pulpy, as in Morning Glory and Mallow, but when dry it becomes fleshy or harder : cnnn-mts, when of the texture of horn, as in codec and the seed of ( 'aiilopliylliun ; and even finny, as in the vegetable ivory, the seed of Phytelephas. It is mostly uniform : but in the nutmeg, FIG. G79. Longitinlmal ma<;ni1ieiT: showing the largo episiierm, the small endosperm in the persistent embryo-sac, and in this the minute- embryo. FIG fi£0. Longitudinal section of ;\ seed of the so-called Pajiaw, Asimina triloba, with ruminated albumen and minute embryo. THE EMBRYO. 311 in the seeds of Asimina (Fig. 680) and all the Custard- Apple family, it is marked by transverse lines or divisions (caused by inflexions or growths of the inner seed-coat) , giving a section of it either a marbled appearance, or as if it had been slit by incisions : it is then said to be ruminated. 5 c 681 (ISJ 684 606. The Embryo,1 being an initial plantlet or individual of a new generation, is of course the most important part of the seed. To its production, protection, and support, all the other parts of the fruit and flower are subservient. 607. In an embryo of full development, namely, one in which all the parts are manifest antecedent to germination, these parts are the Caulicle, otherwise called Radicle, the Cotyledons, and the Plumule. (20, 30.) The first is the initial axis or stem, a primary internode ; the second consists of the leaves of the primary node ; the third is a beginning of a farther growth which is to develop more stem and leaves. Such an embryo is usually unaccompanied by albumen, having in the course of its growth taken into itself (mostly into the cot}'ledons) the provision which in other seeds is mainly accumulated external to it until it is drawn upon in germination. 1 The word Embryo or Embryon was applied to this body in plants by Bonnet (Considerations sur les Corps organise'es),in 1762, and was introduced into systematic botany at about the same time (1763) by Adanson : it was taken up by Gsertner in 1788. Jussieu in the Genera Plantarum (1789) held to the term Cormlnm (the cor semints) which came down from Csesalpinus. Being the germinal part of the seed, the embryo of the plant, like that of the animal, is in general language often called the Germ. FIG. 681. Seed of a Violet (anatropousl, enlarged; with liilum or scar (a), rhaphe (6), and clialaza (c) indicated. 682. Vertical section of the same, showing the straight embryo in the axis of the mealy albumen. FIG. 6S3. Vertical section of the (orthotropous) seed of Buckwheat, showing the embryo folded round in the mealy albumen. FIG. 6S4. Vertical section of the (anatropous) seed of Elodea Virginica, the embryo completely filling the coats FIG. 685. Seed of Delphinium tricorne (anatropous), enlarged; the hilum, the rhaphe. and the chalaza lettered as in Fig. 681. 686. Vertical section of the same with c, the chalaza, d, the testa, e, the tegmen,/, the albumen, g, the minute embryo near the hilum (a). FIG. 687. Embryo of the Pumpkin, with its short radicle and large and flat cotyle- dons, seen flatwise. 688. A vertical section of the same, viewed edgewise. o!2 THE SEED. 608. The opposite extreme is an embryo (as in Fig. C8G) which appears as a mere speck in tin- albumen, lint in which close microscopical inspection may commonly reveal some differ- entiation, such as a slight notch at one end (that farthest re- moved from the micropyle ) ol'a dicotyledonous embryo, indicating the future cotyledons. Indeed, in Monotropese, < )robanchaci •;!•. and some other parasitic dicotyledonous plants, and in Orchids among the monocotyledonous, the embiyo is a globular or oblong particle, with no adumbration of organs whatever antecedent to germination. There are all grades between the most rudimen- tary and the most developed embryos. 609. Under the circumstances of its formation (532), the radicnlar end of the embryo is always near to and points towards the micropyle of the seed, viz. to what was the orifice of the ovule ; and if the embryo be straight, or merely partakes of the curvature of the seed, the cotyledons point to the opposite extremity, that is, to the chalaza. 610. The position of the radicle as respects the hilum varies with the different kinds of seed. In the orthotropous form, as in Ilelianthemum (Fig. 604) and Pepper (Fig. 679), the radicle necessarily points directly away from the hilum.1 In the anatro- pous form, as in Fig. 663, 682, and 684-686, the extremity of the radicle is brought to the immediate vicinity of the hilum ; and so it is, although in a ditl'erent way, in the campy lot ropous seed (Fig. 689, 690) ; while in the amphitropous the radicle points away from the hilum later- ess BOO all}-. As the nature of the ovule and seed may usually be ascertained by external inspection, so the situation 1 Two technical terms, early introduced by Richard to indicate the direc- tion of the radicle (cauliclc), or rather its relation to the hilum, are Aiititrii/ioiis, when the embryo directs its radicle away from the hilum, as it must in nil orthotropous seeds; Orthotropous, also honiotropons, when directed to the hilum (more strictly to the micropylc dose to the hilum), as in anatropmis sci ds. These two terms ;irc still employed by many botanists, although superfluous when the ovule or seed is stated to be anntropous or orthotropous, &c. And the term orthotropous, so used, is liable to be confused with orthotropous as applied to the ovule. Kidianl, moreover, termed the embryo amphitropous when curved or coiled, as in Chick weed (Kin. 080) and all such campylotropous seeds; and In t> m- irojHui^ when neither radicle nor cotyledons point to the hilum, as occurs in the semi-anatropous or amphitropous ovule. Many botanists describe the last by the expression "radicle vague," or, better, " embryo transverse." FIG. C89. C:iiii|iyliitri>)niiis seed of common Cliickweed, magnified- 690. Section ol the same, showing the embryo coiled iuto a riug around the albuinon. THE EMBRYO. 313 of the embryo within, and of its parts, may often be inferred without dissection. But the dissection of seeds is not generally difficult. 611. The direction of the radicle with respect to the pericarp is also noticed by S3'stematic writers ; who emplo}^ the terms radicle superior or ascending when this points to the apex of the fruit ; radicle inferior or descending when it points to its base ; centripetal, when turned toward the axis of the fruit ; centrifugal (or peritropous) , when turned toward the sides ; and vague, when it bears no evident or uniform relation of the kind to the pericarp. 612. The position of the embryo as respects the albumen, when that is present, is various. Although more commonly in the axis, it is often excentric, or even external to the albumen, as in all Grasses and cereal Grains (Fig. 56-61), in Polygonum, &c. When external or nearly so, and curved circularly around the albumen, as in Chickweed (Fig. 690) and Mirabilis (Fig. 17), it is said to be peripheric. 613. The embryo may be very variously folded or coiled in the seed. The two cotyledons, instead of plane and straight, ma}" be crumpled ; or they ma}' be simply convolute or rolled up from one edge, as in Calycanthus (Fig. 691) ; or circinately con- 6l»l 692 693 694 69S volute from the apex, as in Bunias ; or else doubled up and thus biplicately convolute, as in Sugar Maple, Fig. 2. Two modi- fications are more common, and are of such classificatory impor- tance in Cruciferse as to need special reference. Namely, when cotj'ledons are Incumbent (as in Fig. 692, 693), being so folded that the back of one is laid against the side of the radicle ; and Accumbent (Fig. 694, 695),, when the edges of the pair of cotyledons are longitudinally applied to the radicle. These differences were first employed in the classification Cruciferse by FIG. 691. Convolute embryo of Calycanthus, the upper half cut away. FIG. 692. Seed of a Cruciferous plant (Sisymbriuin), with incumbent cotyledons, divided. 693. Embryo of the s;ime detached entire. FIG. 694. Seed of a Cruciferous plant (Barbarea) with accumbent cotyledons. 695. The embryo entire. 314 THE SEED. Robert Brown, and were adopted as primary and tribal characters by DeCandolle. 614. As to number of cotyledons, the two types of embryo are the Monocotyledonous, with a single cotyledon, i. e. leaves at the first nodes alternate (39) ; and Dicotyledonous, with a pair of cot3-ledons, i. e. leaves of the first node in the most simple whorl, a pair, in other words, oppo site (21) ; with its modification of Polycotyledonous (38), the leaves of the first node in whorls of three, four, or more. This occurs with constancy in a majority of ('onii'era* (Fig. 48, 49), occasionally and abnormally in sundry ordinary dicotyledonous species. 615. There are several embryos of the cotyledonous type in which one cot}'ledon is smaller than the other, viz. the inner one when the embiyo is coiled or folded. And in all the species of Abronia (a genus allied to Mirabilis, Fig. 18) this cotyledon is wanting, so that the embiyo becomes technically monocotyle- donous. In another genus, the Dodder (Fig. 78, 79), both cotyledons are constantly wanting ; and the plumule shows only minute scales, the homologues of succeeding leaves reduced almost to nothing. 616. Sometimes the two cotyledons are consolidated into one body by the coalescence of their contiguous faces ; when they are said to be conferruminate. This occurs more or less in the Horsechestnut and Buckeye (Fig. 41, 42), and is striking in the seed of the Live Oak, Quercus virens. 617. The general morphology of the embryo and its develop- ment in germination wen- described at the commencement of this volume. And so the completion of this account of plant, flower, fruit, seed, and embryo brings the history round to the starting point. (12-19, &c.) Having mastered the morphology and general structure of the higher grade of plants, the pupil may go on to the morphology and structure of cells (or Vegetable Anatomy or Histology), and to the study of Cryptogamous Plants in all their grades. TAXONOMY. 315 CHAPTER IX. TAXONOMY. SECTION I. THE PRINCIPLES OF CLASSIFICATION IN NATURAL HISTORY. 618. TAXONOMY, from two Greek words which signify arrange- ment and law, is the study of classification. This is of utmost importance in Natural Histoiy, on account of the vast number of kinds to be set in order, and of relations (of agreement and difference) to be noted. Botanical classification, when complete and correct, will be an epitome of our knowledge of plants. Arrangement according to kinds, and of special kinds under the more general, is common to all subjects of stud}'. But the classification in Biological Natural History, that is in Botany and Zoology, has a foundation of its own. 619. The peculiarity of plants and animals is that they exist as individuals, propagating their like from generation to genera- tion in a series. Of such series of individuals there are very many kinds, and the kinds have extremely various and unequal degrees of resemblance. There are various gradations, but not all gradations of resemblance. Between some, the difference is so wide that it can be said only that they belong to the same kingdom ; between others, the resemblance is so close that it may be questioned whether or not they carne from common parents or near ancestors. 620. The recognition of the perennial succession of similar individuals gives the idea of SPECIES. The recognition of un- equal degrees of likeness among the species is the foundation of GENERA, ORDERS, CLASSES, and other groups of species. 621. Individuals are the units of the series which constitute species. The idea of individuality which we recognize through- out the animal and vegetable kingdoms is derived from ourselves, conscious individuals, and from our corporeal structure and that of the higher brute animals. This structure is a whole, from which no part can be abstracted without mutilation. Each individual is an independent organism, of which the component parts are reciprocally means and ends. Individuality is a main 316 TAXONOMY. distinction between beings and tilings; but. although the tend- ency to individuation begins with life iisrlf. it is conipK-U-l\ realized only in the higher animals. G22. In plants, as also in some of the lower animals, individu- ality is merged in community. No plant ^except one reduced to the simplicity of a single cell, of circumscribed growth, and without organs) is an individual in the sense that a man or a dog is. (16, 15(5.) The herb, shrub, and tree are neither indivisible nor of definite limitation. Whether their successive growths are to remain parts of the previous plant, or to be inde- pendent plants, depends upon circumstances ; and there is no known limit to budding propagation. ('>•>'•>. There is, however, a kind of social or corporate indi- viduality in those animals, or communities (whichever we call them) of the lower grade which are multiplied by buds or oil- shoots as well as by ova, and in which the offspring remains, or may remain, organically connected with the stock. The poly- pidom or polyparium commonly has a certain limitation ami a definite form ; and certain polyps may become organs with special functions subordinate to the common weal. This is more largely true in the vegetable kingdom. So that for de- scriptive purposes, and in a just although somewhat loose sense, the herb, shrub, or tree is taken as an individual. But only while it forms one connected body. Offshoots when separately established are equally individuals in this sense. C>2\. What it is in plants which philosophically answers to the individual in the higher animals is another question, to which various answers have been given.1 Some insist that the whole vegetative product of one seed makes one individual, whether connected or separated (as may happen) into a million of plants. But a common and less strained view restricts the individual to such product only while organically united Others (of which Thouars at the beginning and Braun at the middle of the present century arc1 leading examples) take each axis or shoot with its foliage to represent the individual, of which the leaves and their homologues are organs, the branches 1 icing usually implanted upon the parent axis as this is implanted in the soil, but also equally capable of producing roots by which they may make their own connection with the soil. Still others, on pre- 1 For the history of opinion upon and a full presentation of this topic, see Alexander Braun's Memoir (originally published in the Ahhandl. Akad. Wissenschaften zu Berlin, 1853), Das Individuum der Pflan/.c. &c., and a translation by C. F. Stone in Amer. Jour. Sci. ser. 2, xix. xx is.V. THE PRINCIPLES OF CLASSIFICATION. 317 cisely similar grounds, carry the analysis a step farther, and regard each phytomer (1G) as the individual. Finally, some, in view of their potentially independent life, take the cells, or units of anatomical structure, to be the true individuals ; and this with sufficient reason as regards the simplest ciyptogamous plants. Upon the view here adopted, that plants do not rise high enough in the scale of being to reach true individuality, the question is not whether it is the cell, the phytomer, the shoot, the tree, or the whole vegetative product of a seed which answers to the animal individual, but only which is most analogous to it. In our view, its analogue is the cell in the lowest grades of vege- table life, the phytomer in the higher.1 But, in botanical de- scription and classification, by the individual is meant the herb, shrub, or tree, unless otherwise specified. 625. Species in biological natural history is a chain or series of organisms of which the links or component individuals are parent and offspring. Objectively, a species is the totality oi' beings which have come from one stock, in virtue of that moU; general fact that likeness is transmitted from parent to progeny. Among the many definitions, that of A. L. Jussieu is one of the briefest and best, since it expresses the fundamental conception of a species, i. e. the perennial succession of similar individuals perpetuated by generation. 626. The two elements of species are : 1, community of origin ; and, 2, similarity of the component individuals. But the degree of similarity is variable, and the fact of genetic relationship can seldom be established by observation or historical evidence. It is from the likeness that the naturalist ordinarily decides that such and such individuals belong to one species. Still the like- ness is a consequence of the genetic relationship ; so that the latter is the real foundation of species. 1 For just as successive branches are repetitions and progeny of the parent branch or stem, the phytomers of the branch are repetitions and progeny each of the preceding one, so forming a series of vegetative generations ; and the whole tree might almost as well represent the individ- ual as one of its branches. The phytomer, as well as the branch, is capable of completing itself by producing roots, but is itself indivisible except by mutilation. Least tenable of all is the conception that the whole product of a seed may be taken to represent the vegetable individual. For then individuals increased by buds and division are wholly unlimited both in ex- tent and in duration, so far as observation can show, and a multitudinous race, not only of the present and past, but perhaps in perpetuity, may con- sist of a single individual. There are, indeed, theoretical reasons for infer- ring that a bud-propagated race may not last so long as a seed-propagated species ; but there is no proof of it. See Darwiniana, Art. xii. 318 TAXONOMY. 627. No two individuals are exactly alike ; and offspring of the same stock may differ (or in their progen}- may come to differ) strikingly in SOUK- particulars. So two or more forms which would have been regarded as wholly distinct arc sometimes proved to be of one species by evidence of their common origin, or more commonly are inferred to be so from the observation of a series of intermediate forms which bridge over the differences. Only observation can inform us how much difference is compat- ible with a common origin. The general result of observation is that plants and animals breed true from generation to gem-ra- tion within certain somewhat indeterminate limits of variation; that those individuals which resemble each other within such limits interbreed freely, while those with wider differences do not. Hence, on the one hand, the naturalist recognizes Varieties or differences within the species, and on the other Genera and other superior associations, indicative of remoter relationship of the species themselves. 62S. Varieties are forms of species marked by characters of less fixity or importance than are the species themselves. They may be of all grades of difference from the slightest to the most notable: they abound in free nature, but assume particular importance under domestication and cultivation ; under which variations are most prone to originate, and desirable ones are [in-served, led on to further development, and relatively lixed. 621). If two seeds from the same pod are sown in different soils, and submitted to different conditions as respects heat, light, and moisture, the plants thai spring from them will show marks of this different treatment in their appearance. Such differences an- continually arising in the natural course of things, and to produce and increase them artificially is one of the objects of cultivation. Striking as they often are (especially in annuals and biennials), they are of small scientific consequence. When s|MHii:ineous they are transient, the plant either outlasting the modifying cause or else succumbing to its continued and graver operation. But, in the more marked varieties which alone de- serve (he name, the cause is occult and constitutional; the deviation occurs we know not why, and continues throughout the existence and growth of the herb, shrub, or tree, and con- sequently through all that proceeds from it by propagation from buds, as by offsets, layers, cuttings, grafts, itc. 630. Some varieties of cultivation originate in comparatively slight deviations from the type, and an- led on to greater differ- ences by strict selection of the most marked individuals to breed from. Most appear as it were full-fledged, except as to THE PRINCIPLES OF CLASSIFICATION. 319 luxuriance or development, more or less under the control of conditions, their origin being wholly unaccountable. They arise in the seed-bed, or sometimes from buds, which as the gardeners say " sport." l That is, some seedlings, or some shoots, are unlike the rest in certain particulars.'2 631. Most varieties originate in the seed, and therefore the foundation for them, whatever it may be, is laid in sexual repro- duction. But Bud-variation, or the " sporting" of certain buds into characters in branch, flower, or fruit unlike those of the stock, is known in a good number of plants.3 It might also occur in corals, hydras, and other compound animals propagated by budding. Once originated, these varieties mostly persist, like seedling varieties, through all the generations of budding growth, but are not transmitted to the seed. 632. Upon the general principle that progeny inherits or tends to inherit the whole character of the parent, all varieties must have a tendency to be reproduced by seed. But the inheritance of the new features of the immediate parent will commonly be overborne by atavism, i. e. the tendency to inherit from grand- parents, great-grand-parents, &c. Atavism, acting through a long line of ancestry, is generally more powerful than the heredity of a single generation. But when the offspring does inherit the peculiarities of the immediate parent, or a part of them, its off- spring has a redoubled tendency to do the same, and the next generation still more ; for the tendency to be like parent, grand- parent, and great-grand-parent now all conspire to this result and overpower the influence of remoter ancestry. Close-breed- ing (398) is requisite to this result. In the natural wild state, varieties — many and conspicuous as they often are — must be much repressed b}' the prevalent cross-fertilization which takes place among the individuals of almost all species. Cultivators and breeders in fixing varieties are careful to secure close breeding as far as this is possible. This has fixed the particular sorts of Indian Corn, R}'e, Cabbage, Lettuce, Radishes, Peas, &c., and 1 Both the technical English term. Sport, and its Latin equivalent, are sometimes used for bud-variation only, yet as commonly for seedling variation also. 2 Darwin assumes that variation is of itself indefinite or vague, tending in no particular direction, but that direction is wholly given by the elimina- tion in the struggle for life of all but the fittest for the conditions. But what we observe in the seed-bed does not suggest this view. Ncegeli, Braun, and myself incline to the opinion that each plant has an inherent tendency to variation in certain general directions. 3 A list of known bud-varieties is given in Darwin's Variation of Animals and Plants under Domestication, Chapter xi. 320 TAXn.NM.MY. indeed of nearlv all our varieties of cultivated annual and biennial t/ esculent plants, as well as of several perennials, many of which have lieen fixed through centuries of domestication, while otliers are of recent establishment. What is now taking place with the Teach in this country may convince us that heritable varieties mav l>e developed in trees as well as in herbs, and in the same manner; and that the reason why most races are annuals or biennials is because these can be perpetuated in no other way. and because the desired result is obtainable in fewer year- than in shrubs or trees. Varieties of this fixity of character are called <'>:'•.'!. Races (Lat. I'rolcs). A race, in this technical sense of the term, is a variety which is perpetuated with considerable certainty by sexual propagation. This distinction of varieties pertains chiefly to botany. In the animal kingdom all permanent varieties must be races. So are all indigenous varieties of plants.1 In most of these, the position of species and variet}' is more or less arbitrary or accidental, and capable of interchange. What is called the species may be only a commoner or bettcr- kuown form, or the one first recognized and named by botanists ; whence the other forms as they come to be recognized an1 made to rank in the books as varieties. Instead of one varying from the other, all the forms have probably varied ages ago from a common t3'pe. ('>."• 1. These varieties of the highest order and most marked characteristics, being perpetuable by seed, have the principal attributes of species. They are a kind of subordinate derivative species. Hence they arc sometimes called £< /•;<>*. We judge them not to be so many species, either because in the case of cultivated races we know something of their origin or history. and more of the grave changes which long domestication may bring to pass; or because the forms, however stable, differ among themselves less than recognized species generally do; 01 because very striking differences in the extremes are connected by intermediate forms. And our conclusions, it must be under- stood, "are not. facts, but judgments, and largely fallible judg- ments." For while some varieties appear strikingly different, some species are very much alike.3 1 The Horseradish and a few other plants of spontaneous Lrrowth, which through loim dependence on hud-propagation seem to have lost the power of setting seed, can hardly be called varieties. 2 Darwiniana, 35. :! Wherefore, since we hardly need the term raee in the restricted sense of seed-propagated variety, it is sometimes convenient to use it in the niJin- ner proposed by Uuntham (Anniversary Address to the Linnean Society, THE PRINCIPLES OF CLASSIFICATION. 3*21 635. One distinction between varieties and species is note- worthy and important, even if it may not serve as a criterion. The individuals of different varieties in plants interbreed as freely as do those of the same variety and are equally prolific. Their union produces 636. Cross-breeds.1 In nature, cross-breeding doubtless re- presses variation or prevents the segregation of varieties into what would be ranked as species. In cultivation and domesti- cation, it is turned to important account in producing intermediate new varieties (cross-breeds) variously combining the different excellencies of two parent individuals or two varieties. Thus the great number of forms produced b}' variation (especially as to flowers and fruits) have been further diversified, and selected forms improved for special uses by judicious combination. 637. In general, the individuals of distinct species do not interbreed, although many are capable of it. There is great diversity in this regard among plants, some (such as Willows, Verbascums, and Verbenas) interbreeding freely and reciprocally ; some interbreeding in one direction, but not reciprocally ; others, even when veiy similar, refusing to unite. But, on the whole, there seems to be few nearby related species in which the pollen of the one cannot be made to act upon the ovules of the other by persistent and proper management. Such crossing is an important resource in horticulture. Crossing of species, when successful, produces 638. Hybrids. In these, the characteristics of the two species are combined, sometimes in equal proportions, sometimes with great preponderance of one or the other parent ; and there is often a difference in the result in reciprocal fertilizations. Hy- brids do not play a very prominent part in nature, apart from cultivation, although the limits of some species ma}' be obscured by them, possibly of more than is generally supposed. In the animal kingdom, all the most familiar hybrids are sterile : in the vegetable kingdom, a majority may have a certain but very low degree of fertility ; but this is also the case in many unions May, 1869, 5) as the common designation of any group or collection of indi- viduals whose characters are continued through successive generations, whether it be permanent variety, subspecies, species, or group consisting of very similar species, the term not implying any decision of this question. If this use of the term race prevails, Subsfierii'x will probably take its place as the designation of the highest grade of variety. The objection to this is that the subspecific and specific names would be more liable to be confused. 1 Half-breed is a common equivalent term in the animal kingdom : Latin, Mistus or Mixtus ; French, Metis. 2i 322 TAXONOMY. within the species, and especially in the application of the pollen to the stigma of the- same blossom. Commonly the sterility of hybrids is owing to the impotence of the stamens, which perfect no pollen : and most such hybrids may be fertilized by the pollen of the one or the other parent. Then the offspring either in the first or second generation reverts to the fertilizing species. Moreover, certain hybrids, such as those of Datura, which arc fully fertile per se, divide in the oll'spring, partly in the lir-t gen- eration, and completely in two or three succeeding generations, into the two component species, even when close-fertilized.1 (In part this may come from ad ventive embryo-formation, .">:;:;.) !). There appears, therefore, to be a real ground in nature for species, notwithstanding the difficulty and even impossibility in many cases of defining and limiting them. 640. Species is taken as the unit in zoological and botanical classification. Important as varieties are in some respects, especially under domestication and cultivation, they figure in scientific arrangement only as fractions of species. Species are the true subjects of classification. The aim of systematic natural history is to express their relationship to each other. 641. The whole ground in nature for the classification of spe- cies is the obvious fact that specie's resemble or differ from each other unequally and in extremely various degrees. If this were not so, if related species differed one from another by a constant quantity, so that, when arranged according to their resemblances, the first differed from the second about as much as the second from the third, and the third from the fourth, and so on, -- or if the species blended as do the colors of the rainbow. — then, with all the diversity in the vegetable kingdom then- actually is, there could be no natural foundation for their classification. The mul- titude of species would render it necessary to classify them, but the classification would be wholly artificial and arbitrary. The actual constitution of the vegetable kingdom, however, as ap- pears from observation, is that some species resemble each other very closely indeed, others differ as widely as possible, and be- tween these the most numerous and the most various grades of 1 According to Naudin in Oomptes Kcndus, xlix. is.v.i. & lv. 1862. See also Xauilin's memoir on hybridity in plants in Ann. Sri. Nat. ser. 4, xix. ISC,:!, pp. 180-20:], ^ in Mem. ACM.]. Sri. . . . For tin- literatim' on vegetable hybrid.-, see Ku'lreuter, Xaclirirht, &(•., 17(11, and Appendices, 17(5:5-17(50; Herbert, on Ainaryllid:ieea', 18o~ ; ('. F. (ijrrtner, Versiirlie und Beobachtun- gen ueber die Bastarderzeugung in 1'rlan/enrcieh, 1840; Wirbura, Die Bastardbefrnchtung im Pllan/enreirh, crliiutertert an den Bastarden der Weiden ; and the memoir of Naudin referred to. THE PRINCIPLES OF CLASSIFICATION. resemblance or difference are presented, but always with a mani- fest tendency to compose groups or associations of resembling species, — groups the more numerous and apparently the less definite in proportion to the number and the nearness of the points of resemblance. These various associations the naturalist endeavors to express, as far as is necessary or practicable, by a series of generalizations, the lower or particular included in the higher or more comprehensive. All kinds of differences are taken into account, but only the most constant and definite ones are relied on for characters, i. e. distinguishing marks. Linnaeus and the naturalists of his day used names for only three grades of association, or groups superior to species, viz. the Genus, the Order, and the Glass ; and these are still the principal members of classification. 642. Genera (plural of Genus) are the more particular or special groups of related species. They are groups of species which are much alike in all or most respects, - - which are con- structed, so to say, upon the same particular model, with only circumstantial differences in the details. They are not neces- sarily nor generally the lowest definable groups of species, but •tre the lowest most clearly definable groups which the botanist recognizes and accounts worthy to bear the generic name ; for the name of the genus with that of the species added to it is the scientific appellation of the plant or animal. Constituted as the vegetable and animal kingdoms are, the recognition of genera, or groups of kindred species, is as natural an operation of the mind as is the conception of species from the association of like indi- viduals. This is because many genera are so strongly marked, at least so far as ordinary observation extends. Every one knows the Rose genus, composed of the various species of Roses and Sweetbriers ; the Bramble genus, comprising Raspberries, Blackberries, &c., is popularly distinguished to a certain extent ; the Oak genus is distinguished from the Chestnut and the Beech genus ; each is a group of species whose mutual resemblance is greater than that of any one of them to any other plants. The number of species in such a group is immaterial, and in fact is very diverse. A genus may be represented by a single known species, when its peculiarities are equivalent in degree to those which characterize other genera. This case often occurs ; al- though, if this were universally so, genus and species would bo equivalent terms. If onl_y one species of Oak were known, the Oak genus would have been as explicitly discerned as it is now that the species amount to three hundred ; and better defined, for now there are forms quite intermediate between Oak and 324 TAXONOMY. Chestnut. Familiar illustrations of genera in the animal king- dom are furnished by the (at kind, to which belong the domestic Cat, the Catamount, the Panther, the Lion, the Tiger, the Leop- ard, &c. ; and by the Dog kind, which includes with the Dog the different species of Foxes and Wolves, the .lackal, &c. The languages of the most barbarous as well as of civilixed people everywhere show that they have recognized such groups. Natu- ralists merely give to them a greater degree of precision, and indicate what the points of agreement are. lil.'J. If most genera were as conspicuously marked as those from which these illustrations are taken, genus would be as defi- nitely grounded in nature as species. But popularly recognized genera, rightly based, are comparatively few. Popular nomen- clature, embodying the common ideas of people, merely shows that generic groups are recognizable in a considerable number of cases, but not that the whole vegetable or the whole animal kingdom is divisible into a definite number of such groups of equally or somewhat equally related species. The naturalist discerns the ground of genera in characters which the casual and ordinary observer overlooks ; and, taking the idea of genera from the numerous well-marked instances as the norm, applies it as well as possible to the less obvious or less natural cases, and groups all known species under genera. Resemblances among the species when rightly grouped into genera, though real, are often so unequal in degree, that certain species ma}' be about as nearly related to neighboring genera. So that the recognition of genera even more than of species is a matter of judgment, and even of conventional agreement as to how and where a certain genus shall be limited, and what particular association of species shall hold the position of genus. All the species of a genus must accord in every important structure; but extended observation only can settle the question as to what are important and what are incidental characters. For example, tin' pinnatilid or sinu- ate leaf might have been thought as essential to the < >ak genus as the acorn-cup; but many Oaks are now known with entire leaves, resembling those of Willow or Laurel. An open acorn- cup beset with imbricated scales is a character common to all Kuropean and American Oaks ; but in numerous Asiatic species the cup bears concentric or spiral lamella1 instead, and in others the cup takes the form of a naked and closed sac. Maples have palmately-veincd and lobed leaves; but one species has undi- vided and pinnately-veined leaves. The Apple and the Pear under one view are of the same genus, under another they rep- resent different genera. THE PRINCIPLES OF CLASSIFICATION. 325 644. The genus must be based on close relationship of species, but not necessarily on the closest. Raspberries differ from Blackberries, but must be ranked in the same genus ; and so of Plums and Cherries. For the groups which are to bear the generic name must be as distinct and definite as possible. 645. Orders are to genera what genera are to species. They are groups of a higher rank and wider comprehension, expressive of more general resemblances, or, in other language, of remoter relationship. As all species must be ranked in genera, so all genera must be ranked in orders. FAMILY in botany is Sj'nony- mous with order : at least natural orders and families (however distinguished in zoolog}') have always in botany been inter- changeable terms, and will probably so continue.1 646. As examples of orders in the vegetable kingdom take the Oak family, composed of Oaks, Chestnuts, Beeches, &c. ; the Pine family, of Pines, Spruces, Larches, Cedars, Araucaria, Cypresses, and their allies ; the Rose family, in which Brambles, Strawberries, Plums and Cherries, Apples and Pears are asso- ciated with the Rose in one somewhat multifarious order. 647. Classes are to orders what these are to genera. They express still more comprehensive relations of species ; each class embracing all those species which are framed upon the same broad plan of structure, however differently that plan may be carried out in particulars. 648. Kiiigdom must be added, to represent the highest gener- alization. All subjects of biological classification belong either to the vegetable or animal kingdom. Mineralogy, Chemistry, &c., ma}' use the same terms (genus, species, &c.) in an analo- gous way ; but the classification of substances rests on other foundations than that of beings. O 649. The sequence of groups, rising from particular to univer- sal, is Species, Genus, Order, Class, Kingdom; or, hi descending from the universal to the particular, KINGDOM, CLASS, ORDER, GENUS, SPECIES. 1 Order is the older term, and that which associates best with the technical Latin names. Family is a happy term, which associates itself well with English names. But its use is attended with this incongruity, that the tribe (653) in natural history classification is subordinate to the family. In zoology, order is distinguished from family as the next higher grade. 326 TAXONOMY. i <;:><). This is the common f mmc work of natural history classi- fication. All plants and all animals belong to some species; every species to some genus; every iienn-> to some order or family; every order to some class; every class to one or the other kingdom.1 But this framework, although all that is re- quisite in some parts of natural history, does not express all the ol»ervalile gradations of relationship among species. And even gradations I >elow species ha\e sometimes to IK- classified. The Aeries is capable of extension ; and extension is often requisite on account of the large number of objects to be arranged, and the various degrees of relationship which may come into view. (',.") 1. This is effected by the intercalation of intermediate grades, to be introduced into the system only when there is occasion for them. And in botany one or more grades superior to the classes are needful ; for first and foremost is the great division of all plants into a higher and a lower SERIES 2 (or sub- kingdom), the I'li;enogainous and the Cryptogamous. 652. The grades intercalated into the long-established sequence of Class, Order, Genus, and Species, with new names, are mainly *wo, Tribe and Cohort. 653. Tribe has been for a generation or two thoroughly estab- lished in both kingdoms, as a grade inferior to order and supe- rior to genus. In botanical classification, much use is made of this grade, genera being grouped into tribes. 654. Cohort (Lat. Cohors) is of more recent introduction, at least in Botamr, but is becoming established for a grade next above that of order. Orders are grouped into cohorts. Lindley hit upon a good English name for this grade, that of Alliance. But this word has no available Latin equivalent: while cohort takes equally well a Latin or an English form. i'>~i~>. Finally, each grade is capable of being doubled by the recognition of one like it and immediately subordinate to it, and with designation directly expressive of the subordination. For 1 Nut reeogni/ing TliBckel's third kingdom of Protista, consisting of those Imvrst forms of bring from which the animal and vrgrtalilr kingdoms emerge. - Answering to the French Embranchement in zoology. For this it is pro- posed to use the won! [ >'irhi<»i (IHrixin) : see Laws of Botanical Nomencla- ture ndmited by the International Botanical Congress held at Paris in August, isdT ; to!_ret her with a Historical Introduction and a Commentary, by Alph. DcCandnlle, — Fn^lish translation, London, 18(18; the original French edition. Paris. 1SC7. Perhaps no better name can be found; but the elder DeCandolle brought ftin'sio into common use for a grade subordinate to tribe. F.ndlicher employed the term Regio. We have used Series, and much prefer it THE PKINCIPLES OF CLASSIFICATION. 327 example, if Dicotyleclones and Monocotyledones be the two classes of Phrenogamia, the former (and only the former) is divided upon very important characters into two branches, of far higher rank than the cohorts, viz. the Angiospermae and the Gymnospermse, which take the name of SUBCLASSES. Orders, especially the more comprehensive ones, often comprise two or more groups so distinct that it may fairly be a question whether they are not of ordinal rank : such take the name of SUBORDERS. Tribes in like manner may comprise groups of similar relative value : these are SUBTRIBES. Genera may comprise sections of species which might almost as well rank as genera themselves : to mark their importance and pretension (which ma}- come to be allowed), they are termed SUBGENERA. Finally, forms which are ranked as varieties, but which may establish a claim to be distinct species, are sometimes termed SUBSPECIES. Even what we regard as a variety may comprise more or less divergent forms, to be distinguished as SUBVARIETIES. 656. Some of the larger and most diversified orders, tribes., genera, or species may require all these analytical appliances, and even more, for their complete elucidation ; while others, comparatively homogeneous, offer no ground for them. But when these grades, or some of them, come into use, they are alwaj's in the following sequence : — KINGDOM, SERIES or Division, or Sub-kingdom, CLASS, Subclass, Cobort, ORDER or Family, Suborder, TRIBE, Subtribe, GENUS, Subgenus, Section, Subsection, SPECIES, Subspecies or Race, Variety, Subvnriety. 657. Nature and Meaning of Affinity. These grades, the higher including the lower, denote degrees of likeness or difference. Plants belonging respectively to the two great series or primary divisions may accord only in the most general respects, in that which makes them plants rather than animals. Plants of the same variety are generally as much alike as if they were of the 328 TAXONOMY. same immediate parentage. All plants of the same species are so much alike that they are inferred to have descended from a common stock, and their differences. h»\\c\er grave, are sup- posed to have arisen from sul»eqiient variation, -and the more marked diU'ereiiees to have become lixed through lieredity. This is included in llie idea of species. Descent from a common niiniii explain^ the likeness, and is the only explanation of it. (i.VS. But what is the explanation of the likeness between the species themselves? As respects nearly related species, the answer is clear. Kxcept for practical purposes and in an arbitrary wav, no certain and unfailing distinction can be- drawn between \arietiesofthe highest grade and -pecies of closest resemblance. Il cannot reasonably be doubted that they are of similar origi- nation. Then there are all gradations between very closely and less closely related species of the same genus of plants. C.V.i. The Theory of Descent, that is, of the diversification of the species of a genus through variation in the lapse of time, affords the only natural explanation of their likeness which has yet been conceived. The alternative supposition, that all the existing species and forms were- originally created as they are, and have come down essentially unchanged from the beginning, offers no explanation of the likeness, and even assumes that there is no scientific explanation of it. The hypothesis that the species of a genus have become what they are by diversification through variation is a very old one in botany, and has from time to time been put forward. But until recently it has had little influence upon the science, because no clear idea had been formed of any natural process which might lead to such result. Doubtless, if variation, such as botanists have to recognize within tin- species, be assumed as equally or even more operative through long ante- rior periods, this would account for the diversification of an original species of a genus into several or many forms as differ- ent as those \\hich we recognize as species. Hut this would not account for the limitation of species, which is the usual (but not universal) characteristic, and is an essential part of the idea of species. .lust this is accounted fop by GOO. Natural Selection. This now familiar term, proposed by Charles Darwin, was suggested by the operations of breeders in the development and fixation of races for man's use or fancy ; — in animals by breeding from selected parents, and selecting for breeding in each generation those individuals only in which the desired points are apparent and predominant ; in the seed-bed by rigidlv destroying all plants which do not show some desirable variation, breeding in and in from these, with strict selection of THE PRINCIPLES OF CLASSIFICATION. 329 the most variant form in the particular line or lines, until it be- comes fixed by heredity and as different from the primal stock as the conditions of the case allow. In nature, the analogous selection, through innumerable generations, of the exceedingly small percentage of individuals (as ova or seeds) which ordi- narily are to survive and propagate, is made by competition for food or room, the attacks of animals, the vicissitudes of climate, and in fine by all the manifold conditions to which they are exposed. In the Struggle for Life to which they are thus inevi- tably exposed, only the individuals best adapted to the circum- stances can survive to maturity and propagate their like. This Survival of the Fittest, metaphorically expressed by the phrase natural selection, is in fact the destruction of all weaker com- petitors, or of all which, however they might be favored by other conditions, are not the most favored under the actual circum- stances. But seedlings varying, some in one direction, some in another, are thereby adapted to different conditions, some to one kind of soil or exposure, some to another, thus lessening the com- petition between the two most divergent forms, and favoring their preservation and farther separation, while the intermediate forms perish. Thus an ancestral type would become diversified into races and species. Earlier variation under terrestrial changes and vicissitudes, prolonged and various in geological times since the appearance of the main types of vegetation, and the attendant extinctions, are held to account for genera, tribes, orders, &c., and to explain their actual affinities. Affinity under this view is consanguinity ; and classification, so far as it is natural, ex- presses real relationship. Classes, Orders, Tribes, &c., are the earlier or main and successful branches of the genealogical tree, genera are later branches, species the latest definitely developed ramifications, varieties the developing buds.1 661. Except as to those changes in size, luxuriance, or depau- peration and the like, in which plants, especially seedlings, respond promptly to external influences, as to heat or cold, 1 For the inception of this theory of descent in the form which has within the last twenty years profoundly affected natural history, and developed a copious literature, see a short paper On the Variation of Organic Beings in a State of Nature ; On the Natural Means of Selection ; and On the Com- parison of Domestic Races and True Species, by Charles Darwin, also On the Tendency of Varieties to depart indefinitely from the Original Type, by Alfred Russell Wallace, both read to the Linnean Society, July 1, 1858, and published in its Journal of the Proceedings, iii. (Zoology) 45-62. For the development of the doctrine, see Darwin's " Origin of Species by Means of Natural Selection," "The Variation of Animals and Plants under Domes- tication," and various other works; Wallace's "Geographical Distribution of Animals," £c. For some expositions, see Gray's "Darwiniana." 830 TAXONOMY. moisture or dryness (which are transient and comparatively unimportant), variation, or the unlikene.-s of progeny to parent, is occult and inexplicable. If s< >m< 'limes called out l>v the external conditions, it is by way of internal response to them. In Darwin's conception, variation of itself does not tend in anv Olie particular direction: he appears to attribute all adaptation t<> the sorting which results from the struggle for existence and the survival of the fittest. We have supposed, and N;cgdi takes a similar view,1 that each plant has an internal tendency or pre- di-poMtion to vary in some directions rather than others : from which, under natural selection, the actual differentiations and adaptations have proceeded. I'nder this assumption, and taken as a working hypothesis, the doctrine of the derivation of species serves well for the co-ordination of all the facts in botany, and affords a probable and reasonable answer to a long scries of questions which without it are totally unanswerable. It is sup. polled by vegetable paheontology. which assures us that the plants of the later geological periods are the ancestors of the actual flora of the world. In accordance with it we may explain, in a good degree, the present distribution of species and other groups over the world. It rationally connects the order of the appearance of vegetable types in time with the grades of differ- entiation and complexity, both proceeding from the simpler, or lower and more general, to the higher and more differentiated or special ; it explains by inheritance the existence of function- less parts ; throws light upon the anomalies of parasitic plants in their various gradations, upon the assumption of the most various functions by morphologically identical organs, and indeed illumi- nates the whole field of morphology with which this volume has been occupied. It follows that species are not " simple curiosities of nature," to be catalogued and described merely, but that they have a history, the records of which an1 impressed upon their .structure as well as traceable in their geographical and paheon- lological distribution. This view, moreover, explains the re- markable fact that the characters in which the affinities of plants are mainly discerned (and which therefore serve best for orders, tribes, and other principal groups) are commonly such as are evidently <>f small if any importance to the plant's well- being. and that they run like threads through a scries of species of the greatest diversify in habit, mode of life, and particular adaptations to conditions." t,' mid Begriff dernaturhistorischen Art. Zweite Auflage, 1866. 1 Tfiis is a corollary of natural .selection, which can take ell'ect only U])on useful characters, i. <•. upon structures which play sonic active part BOTANICAL CLASSIFICATION. 331 662. The fixity of species under this view is not absolute and universal, but relative. Not, however, that specific changes are necessitated in virtue of any fixed or all-controlling natural law. Some of the lowest forms have existed essentially unchanged through immense geological periods down to the present time ; some species'even of trees are apparently unchanged in the lapse of time and change of conditions between the later tertiary period and our own day, during which most others have undergone specific modification. Such modifications are too slow to effect in any wise the stability and practical application of botanical classification. SECTION II. BOTANICAL CLASSIFICATION. 663. Natural and Artificial Classifications ma}- be distinguished. A natural classification in botany aims to arrange all known plants into groups in a series of grades according to their resemblances, and their degrees of resemblance, in all respects, so that each species, genus, tribe, order, &c., shall stand next to those which it most resembles in all respects, or rather in the whole plan of structure. For two plants may be very much alike in external appearance, yet very different in their principal structure. Arti- ficial classifications single out one or more points of resemblance or difference and arrange by those, without reference to other considerations, convenience and facility being the controlling principles. The alphabetical arrangement of words in a dic- tionary, and the sexual system in botany b}* Linnaeus (or rather a part of it), — in which plants are arranged in classes upon the number of their stamens, and in orders upon the number of pistils, — are examples of artificial classification. The arrange- ment of the words of a language under their roots, and with the derivative under the more primitive forms, would answer to a natural classification. in the life of the plant, and which therefore undergo modification under (•hanging conditions. Unessential structures accordingly are left unaltered or are only incidentally modified. And so these biologically unessential points of structure, persisting through all adaptive changes, are the clews to relationship. Thus, Rubiaceas are known by insignificant stipules, Ano- nacese by ruminated albumen, Rhamnaceaj by a valvate calyx and stamens before the petals, &c. Paradoxical as it may seem, it is not although, but because, they are of small biological importance that they are of high clas- sificatory (/. r. of genealogical) value. On considerations like these, characters are divided into ada.pt.ire or bio- logical on the one hand, and genealogical or genetic on the other. The saga- cious naturalist seizes upon the latter for orders and the like ; while the former are prominent in genera, &c. 332 TAXONOMY. 664. No artificial classification of plants could fail to be n;itiir:il in sonic portions :iinl some respects ; because plants which a-rcc in any point of structure likely to lie used for the purpose will commonly agree in other and perhaps more impor- tant characters. On the other hand, no natural classification can dispense with artificial helps; nor can it express in lineal order, or in any other way, all the various relationships <,f plants, even if these were full v determined and rightly subordinated. Natu- ralists now endeavor to make classification as natural as possible ; that is, to base it in every grade upon real relationships. What real relationships are. and how to express them in a general s\-tem and throughout its parts, has been the task of the leaders in liotanv from the beginning of the science until now ; and the work is by no means completed. r. < '.."). Linn:eu-> uas perhaps the first botanist to distinguish dearlv between a natural and an artificial classification. lie laliored ineffectually upon a natural classification of the genera of plants into orders ; and he devised an effective artificial classi- fication, which became so popular that it practically superseded all others for more than half a century, and has left a permanent impression upon the science. The last generation of botanists who were trained under it has not quite passed away. r,i;r>. Auto-IJiniivan Classification. Linmi-us, in his Philosophia Ilotanica, divided systematists into heterodox and orthodox: the former, those who classify plants by their roots, herbage, time of flowering, place of growth, medical and economical uses, and the like ; the latter, by the organs of fructification. It is remark- able that all the orthodox or scientific classifications anterior to Linmcus made a primary division of the vegetable kingdom into Trees and Herbs, referring the larger shrubs to the former and the undcr-shrubs to the latter. — an arrangement which began with Theophrastus and was continued by Kay and Tournefort. C.IM. The three most important names in botanical taxonomy anterior to I.iniuvus are those o"Cesalpini. Ray. and Tonrnefort. Scientific botany commenced with the former, in Italy, in the latter half of the sixteenth century. He first used the embryo and its cotyledons in classification, distinguished differences in the in- sertion of floral parts, and. indeed (excepting the primary division into trees and herbs), founded all principal characters upon the organs of fructification, especially upon the fruit and seed. ( onrad (lesner of /urich had somewhat earlier recognized this principle, but Ccsalpini first applied it. 668. A century later ( 1 C'.iO-'.i'.i ) this principle was carried into practice by Kivinus (a name latinized from Bachmann), of BOTANICAL, CLASSIFICATION. 333 Leipsic, in a wholly artificial classification founded on the corolla. His contemporary in England, Robert Morison, somewhat earlier began the publication of his great work, the Universal History of Plants. In this was first attempted a grouping of plants into what are now called natural orders ; and these were defined, some- what loosely, some by their fruit, inflorescence, and flowers, others by their stems, the nature of their juice, &c. But the two great systematists of the time, who together laid the foundations of modern scientific botany, were John Ray in England and Joseph Pitton de Tournefort in France. (i(ii). Ray's method of classification was sketched in 1682, and was anterior to Tournefort's, but was amended and completed in 1703. The leading fault of both was the primary division into trees and herbs. The great merit of Ray was his division of herbs into Flowerless and Flowering, and the latter into Dicotyle- donous and Monocotyledonous. These great classes he divided and subdivided, by characters taken from the organs of fructi- fication, into what we should call natural orders or families, but which he unfortunately called genera. He noted the coincidence of nerved leaves with the monocotyledonous embiyo, although he did not notice that his first division of arborescent plants was monocotyledonous : and he had a clear apprehension of genera. 670. Tournefort's method was published in French in the 3rear 1G94, in Latin in 1700. It is more definite but more arti- ficial than that of Ray, being founded like that of Rivinus almost wholly upon modifications of the corolla, and it overlooked the dis- tinction between monocotyledonous and dicotyledonous embryos. Its great merit is that here genera, as we now understand them, are first established and defined, and all the species then known referred to them ; so that Tournefort was justly said by Linnaeus to be the founder of genera. Ray may be said to have indicated the primary classes, Jussieu (in the next century) to have estab- lished natural orders, and Tournefort to have given to botany the first Genera Plantarum. 671. Linnaean Classification. Linnaeus, the great reformer of botany in the eighteenth century, thoroughly revised the principles of classification, established genera and species upon a more scien- tific basis, and, in designating species by a word instead of a descriptive phrase, introduced binomial nomenclature. (704.) He likewise established for the stamens, and indeed for the pistils also, their supreme importance in classification (probably without knowledge of the clear suggestion to this effect made by Burckhard in a letter to Leibnitz, printed in 1702) ; their functions, so long overlooked, being now ascertained. He also 334 TAXONOMY. drew a clear and practical distinction between natural and arti- ficial classifications ((•(;.'>). ainl deferring all endeavors to make the former available, except for genera. he devised a practical sulistitute for it. as a key to the genera, vi/. his celebrated (\1'2. Sexual System, or arrangement of the genera under arti- ficial classes ami <>nleix, founded upon the stamens and pistils. Although no\v out <>f n>e. this artificial classification has been >o popular and inlliieiit ial. and has left -<> deep an impression upon Hie science and especially upon the language of botany, that it needs ti> lie presented. The primary divisions are the classes, twenty-four in number. 15nt the I' 1th class. Cryptogamia. ccm- si-ts of plants which have not stamens and pistils and conse- quently no proper llowcrs. and is therefore the counterpart of the remaining t \venty-thive classes, to which the corresponding name of rhaneroo-amia or. in shorler funn. I'lia-noo-amia ( 1'ha'no^ainous plants; has since been applied. These twenty-three classes are characterized by certain inodilicatioiis and associations of the slamens. and have substantive names, of (ireek derivation, ex- pre^sive of their character. The first eleven comprise all plants with perfect (/'. r. hermaphrodite) tlowers. and with a definite number of equal and unconnected stamens. They are distin- guished by the absolute number of these organs, and are desig- nated bynames compounded of (ireek numerals and the word . I-'.NM. \M>I:I \, with nine stamens, as in the Rhubarb. 10. l>i.e\\iii:i \. with ten stamens, as in Rhododendron and Kalniia. 11. D«II>I. \\m:i\. \vith twelve stamens, as in Asanim and the Miirno- iii-tie: extended also to include those with from thirteen to nine- teen stamens. U7."i. The two sticceeiliim' classes include plants with ])erfect flowers having twenty or more unconnected stamen-, which, in 12. L «I»\MH:I \. are inserted on the calyx (perigynous), as in the Rose Family ; and in 13. I'm ^ \M.I;I \. <>n the receptacle (hypogynous), as in the Buttercup, Anemone. &c. BOTANICAL CLASSIFICATION. 335 674. Their essential characters are not indicated by their names : the former merely denoting that the stamens are twenty in number; the latter, that they are numerous. - -The two fol- lowing classes depend upon the relative length of the stamens, namely : — 14. DIDYNAMIA, including those with two long and two short stamens, as in the majority of flowers with bilabiate corolla. 15. TETRADYNAMIA, those with four long and two short stamens, as in flowers with cruciferous corolla. 675. These names signify in the former that two stamens, and in the latter that four stamens, are most powerful. - -The four succeeding are founded on the connection of the stamens, viz. : - 16. MONADELPHIA (meaning a single fraternity), with the filaments united in a single set, tube, or column, as in the Mallow. 17. DIADELPHIA (two fraternities), with the filaments united in two sets or parcels, as in Corydalis and in many Leguminosas. 18. POLYADELPHIA (many fraternities), with the filaments united in more than two sets or parcels, as in Hypericum. 19. SYNGENESIA (from Greek words signifying to grow together), with the anthers united in a ring or tube, as in the Sunflower and all Composite. 676. The next class, as its name denotes, is founded on the union of the stamens to the st3'le : - 20. GYNANDRIA, with the stamens and styles consolidated, as in Cypri- pedium and all the Orchis Family. 677. In the three following classes, the stamens and pistils occupj' separate blossoms : - 21. MONCECIA (one household) includes all plants where the stamens and pistils are in separate flowers on the same individual ; as in the Oak and Chestnut. 22. DKECIA (two households), where they occupy separate flowers on different individuals ; as in the Willow, Poplar, Moonseed, &c. 23. POLYGAMIA, where the stamens and pistils are separate in some flowers and associated in others, either on the same or two or three different plants ; as in most Maples. 678. The remaining class is essential!}' flowerless ; or rather its organs of reproduction are more or less analogous to, but not homologous with, stamens and pistils. But, although Liunreus suspected a sexuality in Ferns, Mosses, Algae, &c., there was no proof of it in his day. So he named the class, containing these, 24. CRYPTOGAMIA, meaning clandestine marriage, the sexes, if existent, hidden from view. 679. The characters of the classes may be presented at one view, as in the subjoined table : — TAXONOMY. t- OC C. - — • I -'. O 1-IC3 nler 1. MONOGYNIA, those with one style or sessile stigma to the flower. 2. DIGYNIA, those with two styles or sessile stigmas. 3. TRIGYNIA, those with three styles. 4. TETRAGYNIA, those with four styles. 5. PENTAGYNIA, those with five styles. 6. HEXAGYNIA, those with six styles. 7. HEPTAGYNIA, those with seven styles. 8. OCTOGYNIA, those with eight styles. 9. ENNEAGYNIA, those with nine styles. 10. DECAGYNIA, those with ten styles. 11. DODECAGYNIA, those with eleven or twelve styles. 12. POLYGYNIA, those with more than twelve styles. 681. The orders of class 14, Didynamia, are only two and are founded on the pericarp, namely : - 1. GYMNOSPERMIA, meaning seeds naked, the achenia-like fruits of a 4-parted pericarp having been taken for naked seeds. 2. ANGIOSPERMIA, with the seeds evidently in a seed-vessel or peri- carp, /. e. the pericarp undivided. 682. The 15th class, Tctradynamia, is also divided into two orders, which are distinguished merely by the form of the pod:- 1. SILICDLOSA; the fruit a silicle (561), or short pod. 2. SILIQUOSA; fruit a silique (561), or more or less elongated pod. 683. The orders of the 16th, 17th, 18th, 20th, 21st, and 22d classes depend merely on the number of stamens ; that is, on the characters of the first thirteen classes, whose names they likewise bear: as MONANDRIA, with one stamen, DIANDRIA, with two stamens ; and so on. 684. The orders of the 19th class, Syngenesia, are six, namely : 1. POLYGAMIA /EQUALIS, where the flowers are in heads (the so-called compound flower), and all hermaphrodite. 2. POLYGAMIA SUPERFLUA, the same as the last, except that the rays, or marginal flowers of the head, are pistillate only. 3. POLYGAMIA FRUSTRANEA, those with the marginal flowers neutral, the others perfect. 4 POLYGAMIA NECESSARIA, where the marginal flowers are pistillate and fertile, and the central staminate and sterile. 5. POLYGAMIA SEGREGATA, where each flower of the head [or glom- erule] has its own proper involucre. 6. MONOGAMIA, where solitary flowers (that is, not united into a head) have united anthers, as in Lobelia. 338 TAXONOMY. <;*;"}. The 23d class, Polygamia. has three orders, two of them founded on tin- character of the two preceding classes and bearing their names, and the third named upon the same prin- ciple, namely : - 1. M<>X«KCIA, where both separated and perfect flowers are found in tin- same plant. 2. DKECIA, where they occupy two different plants. 3. TRiuiciA, whciv OIK.' individual bears the perfect, another the stami- na te, and a third the pistillate flowers. 686. The orders of the 24th class, Cryptogamia, the Flower- less Plants, are so many natural orders, and are not definable by a single character. They are : — 1. FILICES, the Ferns. 2. Mcsci, the Mosses. 3. ALG/E, which, as left by Linnaeus, comprised the Hepaticae, Lichens, &c., as well as the seaweeds. 4. FUNGI, Mushrooms, &c. 687. In its day, this artificial system well fulfilled its purpose, and was preferred to all others on the score of facility and defi- niteness. Now no botanist would think of employing it, nor would it be chosen for a key to genera, which was its only legiti- mate use. 688. The Natural System was rightly appreciated by Linnaeus, who pronounced it to be the first and last desideratum in syste- matic botany; and he earl}" attempted to collocate most known genera under natural orders (e. g. Piperitce, Pulmce, Scttammce^ Orchidea, AitK'xtacece, &c., sixty-seven in number, including his four cryptogamic orders), but without definition or arrangement. In his later years, he was unable to accomplish any thing more. The dillieulf problem was taken up by Limueus's contemporary and correspondent, Bernard de .Jussieu. who planted the botanic garden at Trianon with plants grouped into natural orders, but published nolhin-j.-. ITis pupil. Adanson, who when a young man lived for several years in Senegal, and who was as remark- able lor eccentricity as for erudition and ability, published in 17i'.."i. in his Families des Plantes. the first complete system of natural orders. But he seems to have taken little from his teacher, and with all his genius to have contributed little to the advancement of the natural system. 689. Antoine Laurent de Jussieu, nephew of Bernard, has been called the founder of the natural system of botany, and to him more than to any other one person this honor may be ascribed. In his (ienera IMantarum secundum Ordines Xatu- rales disposita, 1789, natural orders of plants, one hundred BOTANICAL CLASSIFICATION. in number, were first established and defined by proper char- acters, and nearly all known genera arranged under them. His primary division of the Vegetable kingdom was into Acotyledones, Munocotyledones, and Dicotyledones, adopted from Ray, with a change which was no improvement. For his Acotyledones, the Cryptogamia of Linnasus, are the '"plants without flowers" of Ra}-; they are, to be sure, destitute of cotyledons (though not in the manner of Cuscuta) , because destitute of embryo altogether. The Acot3'ledones forming his first class, Jussieu divided the Monocotyiedones into three classes upon single and artificial characters, namely upon the insertion of the stamens, whether Irypogynous, perigynous, or epig}'nous ; and the Dicotyledones, into eleven classes on similar characters, preceded b}' a division into Apetalce, Monopetalce, Polypetalce, and Diclines irregulares, i. e. first upon the character of the perianth, then upon the insertion of the stamens or in Monopetalae of the corolla. The following is the scheme : — Acotyledones CLASS I Monocotyiedones Stamens hypogynous II. perigynous III. epigynous IV. Dicotyledones Apetalous . Monopetalous Stamens epigynous ^ V. perigynous VI. hypogynous VII. Corolla hypogynous VIII. perigynous IX. epigynous: anthers connate X. epigynous : anthers separate XI. Stamens epigynous XII. hypogynous .... XIII. perigynous XIV. Polypetalous . Diclinous (also Apetalous) XV. 690. Auguste Pyrame DeCandolle was the next great syste- matist. Reversing the order of Jussieu, who proceeded from the lower or simpler to the higher or more complex forms, DeCan- dolle began with the latter, the phsenogamous or flowering plants, and with those having typically complete flowers. On account of its convenience and the greater facilities for studying the higher plants, this order has been commonly followed ever since. His primary division on anatomical structure, into Vascular and Cellular plants, was a backward step, confusing a portion of the lower series with the higher ; and the duplicate names of Exo- gence and Endogence, appended to Dicotyledoneoe and Monoco- 340 TAXONOMY. tylerlonese, as it now appears should have been omitted. The grades of the Candollcan s \.-tem superior to the orders, in their final form, are mainly these : - Div. I. VASCULAR (more properly PILENOGAMOUS) PLANTS. CLASS I. DICOTYLKDONOUS or EXOGENOUS. Subclass I. THAI.AMIKLOROUS : petals (distinct) and stamens on the torus, /. i .-. free. II. CALYCIFLOROUS : petals (distinct or coalescent) and stamens adnate to the calyx. III. COROLLIFLOKOUS : petals (mostly coalescent) not ad- natu to calyx, bearing the stamens. IV. MONOCHLAMYDEOUS: petals wanting. CLASS II. MOXOCOTYLEDONOUS or ENDOGENOUS. (No subclasses.) Div. II. CELLULAR (more properly CRYPTOGAMOUS) PLANTS. CLASS I. JETIIEOGAMOUS : with sexual apparatus, and Vascular tissue. (AV/'"'*. tacece—FUices.) Only cellular tissue. (Mimri and Hcpaticce.) CLASS II. AMIMIICAMOCS : destitute of sexual organs and of other than cellular tissue. (Lii-ln m.-, l-'mnji, Alijce.) 691. Cryptogaraous plants of all orders are now known to be provided with sexes ; and the Jussiaean divisions of the Dico- tyledones into Apetalce (including Die-lines), Monopetalce, and Polypetala, are generally preferred to those of DeCandolle. Into the present views of the classification of the Cryptogamia it is unnecessary here to enter. Their general arrangement into classes, &c., is not yet well settled, and the whole taxonomy of the lower Cryptogams is at present in a state of transition. (I'.)2. John LindU-y in successive attempts (between is,°,0and 1815) variously modified, and in some few respects improved, the ( 'andollean arrangement. But, as neither his groupings of the natural orders nor the new classes which he adopted have been approved, his schemes need not be here presented. He must be credited, however, with the first attempt to carry into etl'eet a suggestion made by Brown, that the orders should themselves be disposed as far as possible into superior and strictly natural groups. In Lindley's first attempt, such groups of two grades were proposed, the lower called H/.I-HS (tendencies), the higher cohorts. In his later and largest work. The Vegetable King- dom, these were reduced to one. and the name of alliance was coined. But this word lias no good Latin equivalent, and the term cohort (cohors) is preferred. BOTANICAL CLASSIFICATION. 341 693. Robert Brown, next to Jussieu, did more than any other botanist for the proper establishment and correct characterization of natural orders. Having in the year 1827 published his dis- covery of the gymnospermy of Coniferae and C^'cadaceae, it was in Lindley's works that this was first turned to proper systematic account by dividing the class of Dicotyledones into two subclasses, the Angiospennce and the Gymnospermce. The latter has been elevated by the vegetable palaeontologists to the rank of a class. 694. Stephen Ladislaus Endlicher, of Vienna, a contempo- rary of Lindley, of less botanical genius, but of great erudition and aptness for classification, brought out his complete Genera Plantarum secundum Ordines Naturales disposita, between the years 1836 and 1840. This elaborate work follows that of its predecessor, Jussieu, in beginning with the lower series of plants and ending with the higher. Its primary division is into two regions: 1. Thallopliyla, plants without proper axis of growth (developing upward as stern and downward as root), no other tissue than parenchyma, and (as was thought) no proper sexes. This answers to the lower or Amphigamous Cellular plants of DeCandolle. 2. Cormophyta, plants with an axis (stem and root), with foliage, &c. The Cormophyta, or plants of the higher region, Endlicher divided into three great sections: 1. Acro- brya, answering to the higher vEtheogamous Ciyptogamia of DeCandolle, with which was wrongly associated a group of root- parasitic flowering plants (the Rhizantheae) which were fancied to bear spores instead of embryo in their seed ; 2. Amphibrya, which answer to Monocotyledones ; and, '3. Acramphibrya, which answer to Dicotyledons. These last contain five cohorts: 1. Gymnospermece ; 2. Apetalce ; 3. Gamopetalce (the Monopetalne of Jussieu better named) ; 4. Dlali/petalce (the Polypetahe of Jussieu, &c.)- The cohort in Endlicher's classification, it will be seen, is a higher grade than that to which this name was applied by Lindley in the more recent use. For the latter, i. e. for the grade between these and the order, Endlicher employed the name of class. 695. Finally, the Genera Plantarum, now in course of pub- lication by George Bentham and Joseph Dalton Hooker, adopts in a general way the Candollean sequence of orders, with vari- ous emendations ; divides the class of Dicotyledons into two sub- classes, Angiospermous and Gymnospermous ; the former into the Polypetalous, Gamopetalous, and Apetalous divisions ; and the first of these into the Thalamiflorous, Disciflorous, and Caly- ciflorous ' ' series " (the middle one composed of the latter part 342 TAXONOMY. of DeCandolle's Thalamiflora? with some of his Caiycifloroe) ; and under these the orders are arranged in cohorts, — fifteen cohorts in the Poly pet ahe, and ten under three "series" in the Gamopetahe. The remainder of this particular classification has not yet appeared in print, although partly sketched by its authors. It will generally lie adopted in this country, with some occasional minor modifications. r.'.ii;. Various modifications have been from time to time pro- posed. One of the best of them in principle is that initiated by Adolphe Brongniart and adopted by many European botanists, which, recognizing that most apetalous dowers are reductions or degradations of polypetalous types, intercalates the Apctalae or Monochlaniyde;e among the Polypetalae. But this has never yet been done in a satisfactory manner, or without sundering orders which should stand in contiguity. C>'.»7. It should be borne in mind that the natural system of botany is natural only in the constitution of its genera, tribes, orders, &c., and in its grand divisions; that its cohorts and the like are as yet only tentative groupings ; and that the putting together of any or all these parts in a sj-stem, and especially in a lineal order, necessary as a lineal arrangement is, inu-t needs be largely artificial. So that even the best perfected arrangements must always fail to give of themselves more than an imperfect and considerably distorted reflection of the plan of the vegetable kingdom, or even of our knowledge of it.1 1 In the first place, tin- relationships of any group cannot always be rii^litlv estimated before all its members are known and their whole struct- ure understood ; so that the views of botanists are liable to be modified with the discoveries of every year. The discovery of a single plant, or of a point of structure before misunderstood, has sometimes changed materially the position of a considerable group in the system, and minor alterations are continually made by our increasing knowledge. Then the uroups which we recognize, and distinguish as genera, tribes, orders, &c., are not always, and perhaps not generally, completely circumscribed in nature, as we are obliged to assume them to be in our classilication. This might be expected from the nature of the case. For the naturalist's groups, of whatever grade, are not /•'<.< of resent- liliiiKT, which may In' expected to present infinite gradations. Mesides, al- though the grades of allinity among species arc most various, if not wholly indefinite, the naturalist reduces them all to a few, and treats his genera, tribes, &c., as equal units, or as distinguished by characters of about equal value throughout, — which is far from being the case. And in Ids works he is obliged to arrange the groups he recognizes in a lineal series; but each genus or order, >xc . is very often about equally related to three or four others : so that only a part of the relationship of plants can in any way be indicated by a lineal arrangement. BOTANICAL CLASSIFICATION. 343 698. Even the great classes cannot be arranged in a single line, beginning with the highest Phaenogains and leaving the lowest in contiguit}' with the higher Cryptogams. The Dicoty- ledons take precedence of the Monocotyledons in rank. Yet a part of them, the Gymnosperms, are much the lowest of all known Phsenogams as regards simplicity of floral structure ; and through them only is a connection with the higher Cryp- togams to be traced. The Monocotyledons stand upon an iso- lated side line, and have no such simplified representatives. In placing the latter class between the Dicotyledons and the Acrogens, the chain of affinities is widely sundered. If, yield- ing to a recent tendenc}', we raise the Gymnosperms to the rank of a class, and place it between the Monocotyledons and the Acrogens, then the much nearer relationship of Gymno- sperms to Angiosperms through Gnetaceae and Loranthaceae is not respected. (606, &c.) 699. Nor can the angiospermous Dicotyledons be disposed lineally according to rank. The apetalous and achlamydeous must be the lowest. Some are evidently reduced forms of Poly- petalae or even of Gamopetalae : the greater part cannot without violence be thrust into their ranks. The Gamopetalae, especially those with much floral adnation, should represent the highest type, the organs being at the same time complete and most dif- ferentiated from the foliar state. If a natural series could be formed, these would claim the highest place, with the Compositae perhaps at their head. In the Candollean sequence, they occupy the middle ; and the series begins, not without plausible reason, with orders having generally complete blossoms, and such as most freely and obviously manifest the homology of their organs with leaves, then rises to those of greater and greater combi- nation and complexity, and ends with those plants which, with all their known relatives, are most degraded or simplified by abortions and suppressions of parts which are represented in the complete flower. These are low in structure, equally whether we regard them as reduced forms of higher types, or as forms which have never attained the full development and diversifica- tion which distinguish the nobler orders. 700. Actual classifications, in their leading features and in their extension to the cohorts, orders, &c., must be studied in the systematic works where they are brought into use. In these are the applications of the principles which are here outlined. A separate volume of this text-book should illustrate the structure, relations, and most important products of the phaenogamous natural orders, as another is to illustrate the 344 TAXONOMY. ciyptogamous orders. A s}'noptical view of the great divi- sions 01113-, as at present received and named, is appended. Definitions and characters may be sought in the present and preceding chapters. SERIES I. PH^ENOGAMOUS OR FLOWERING PLANTS. CLASS L DICOTYLEDONS. SUBCLA>> I AXGIOSPERMS. l>ir. 1. I'oLYTKTALOUS. l>ir. '1. (i.VMOI'KTALOUS. (MonOpetdloUS.) Div 3. APETALOUS. SUBCLASS II. GYMNOSPERMS. CLASS II. MONOCOTYLEDONS. Div. 1. SPADICEOUS. Div. 2. PETALOIDEODS. Div. 3. GLUMACEOUS. SERIES II. CRYPTOGAMOUS OR FLOWERLESS PLANTS. CLASS III. ACROGENS. Div 1. VASCULAR. (Ferns and their allies.) Div. 2. CELLULAR. (Mosses and Liverworts.) CLASS IV. THALLOGENS OR THALLOPHYTES. PHYTOGRAPHY. 845 CHAPTER X. PHYTOGRAPHY. SECTION I. NOMENCLATURE. 701 . PHYTOGRAPHY is the department of botany which relates to the description of plants. This includes names and terms, also figures and signs, as well as characters and detailed descriptions. It comprises two sorts of names, one used to designate organs or modifications of organs, the other to distinguish plants or groups of plants. The former is Glossology or (to use the more common but less proper word) Terminology. The latter is prop- erly Nomenclature. 702. Names of Plants were at first only generic names. The language of botany being Latin, and the plants which the old herbalists knew being mostty European, their scientific names were mainly adopted from the ancient Romans or, through Latin literature, from the Greeks. Ex. Quercus, Prunus, Rosa, Rubus, Trifoliurn ; and of Latinized Greek names, Agrostis, Aristolochia, Colchicum, Melilotus. To the classical names others were added from time to time ; as, from the Latin, Bidens, Convallaria, Dentaria ; from the Greek, Anacardium, Glycyrrhiza, Loranthus, &c. Some barbarous or outlandish names were early adopted, such as Alhagi from the Arabs, and Adhatoda and Nelumbo from India. These are mostly such as were or could be conformed to Latin ; as Datura and Ribes from the Arabic, and later Thcea and Cojfcea. Of American aboriginal names, ffura, Guaiacum, and Yucca are examples. Some ancient names of plants commem- orated distinguished men. Ex. Asclepias, Euphorbia, Lysimachia, Pceonia. Tournefort and his contemporaries resumed this practice, and named plants in memoiy or in honor of distin- guished botanists. Ex. Begonia, Bignonia, Ccesalpinia, Fuchsia, Gerardin, Lobelia, Lonicera, Magnolia. 703. When among plants of the same name or kind different species were known, these were distinguished by annexed epi- thets. For example, among the Pines there were : Pinus syl- vestris, vulgaris ; Pinus sylvestris, montana altera ; Pinus sylvestris, montana tertia ,' Pinus sylvestris maritima, conis firmiter ramis adhcerentibus ; Pinus maritima minor ; Pinus maritima altera, &c. 346 PHYTOGRAPHY. And as the number of known species increased, so did the length of the phrases which were needed for their discrimina- tion. These " differentiae," thus used as specific names (the nomina specified of Linnaeus), became extremely cumbrous. It was about in the middle of his career that Linnaeus suggested what he called tririul names (nomina trivialia) for the specific name, consisting of a single word ; and in the Species Plan- taruin, in 1753, he carried this idea into full effect in Botany. The step was a simple one, but most important; and LinnuHis himself, who generally did not underrate his services to science, seems hardly to have appreeiated its practical value.1 704. The Binomial Nomenclature in Natural History, thus established, first separated the name of a plant or of an animal from its diagnosis, descriptive phrase, or character, and reduced the appellation to two words, the first that of its genus, the sec- ond that of its species. The generic name very nearly answers to the surname of a person, as Brown or Jones ; the specific answers to the baptismal name, as John or James. Thus, Quercus alltu is the botanical appellation of the White Oak ; Quercus being that of the genus, and n (white) that of a particular specie's ; while the Red Oak is named Quercus rubra ; the Scarlet Oak. Quercus cocciitcii; the Live Oak, Quercus virens ; the Ilur Oak, Qm-mis macrocarpa: Magnolia grandiflora is Large-flowered Magnolia; M. inacrophylla, Long-leaved Magnolia, and so on. The name of the genus is a substantive, or at least is a word taken as a suli-tan- ti\e. That of a species is mostly an adjective adjunct, always following the generic name and in the same gender.'2 This com- bination of generic and specific name is the name of the plant.3 70.3. By this system, not only is the name of the plant reduced to two words, but a comparatively moderate number of words serves for the complete designation of more than 120,000 plants,4 1 Moreover, he may lie said to have adopted rather than originated the idea; for single-worded specific names were used half a century previous by Bachmami, u/ins Kivinus. - It is to lie noted that the classical Latin names of trees are all feminine, therefore (tun-nix II//HI, /'inns rii/iiln, &C. 1 The name of a subgcnus is sometime-' written in between the two parts of the plant's name, as /'/•»;)/(.•>• (failns) Vii-i/iiiimxt. This is the name of the plant and something more. In addition to the name of the species, that of the variety or even suhvariety is sometimes added. 4 Alphonsr 1 )c( 'andolle several years ago estimated the known species of Flowering Plants at between HID, unit and r.'O.OOi). The larger number may perhaps include the higher orders of the Flowerless series. In the pre-ent state of our knowledge of the lower orders of Cryptogams, no close estimate can be well formed of the actual number of species. NOMENCLATURE. 347 in a manner which avoids confusion and need not overburden the memory. The generic part of the name is peculiar to each genus. The specific adjunct is not available for more than one species of the same genus, but may be used in any other genus. They are so widely thus employed that the number of specific may not exceed that of generic appellations. 706. To render this sj'stem of nomenclature most serviceable for the ready identification of such numbers of plants or groups, and for the clear and succinct presentation of or reference to what is known and recorded of them, rules are indispensable, and conformity to admitted rules is a manifest duty. Such rules were systematically formulated first by Linnaeus, in his Funda- menta, Critica, and Philosophia Botanica, chiefly for generic names, some of them being of the nature of laws, some rather of recommendations. The most important of them remain in full force, while many of the more particular rules restricting the choice of names have been abandoned. The code was judi- ciously revised (in his Theorie Elementaire) by DeCandolle " who was ruled by the idea of having the law of priori ty prop- erly respected," was critically considered by Lindle}' in his In- troduction to Botany, and has of late been reformulated by Alphonse DeCandolle under the sanction of a Botanical Congress Held at Paris in 1867.1 707. Rules for Naming Plants. These "should neither be arbitrary nor imposed by authority. They must be founded on considerations clear and forcible enough for every one to com- prehend and be disposed to accept. The essential point in nomenclature is to avoid or to reject the use of forms or names 1 Lois de La Nomenclature Botanique, etc., Geneva and Paris, 18fi7. In the English edition, translated by Weddell : Laws of Botanical Nomenclature adopted by the International Botanical Congress held at Paris in August, 1867, together with an Historical Introduction and a Commentary, London, Reeve & Co., 1808. The Laws, simply, were reprinted in the American Journal of Science and Arts, July, 1868. A few special points have been more recently discussed by various critics, especially in the Bulletin of the Botanical Society of France, and in that of the Royal Botanical Society of Belgium. See like- wise American Journal of Science and Arts for September, 1870, and August, 1 S77 ; also, Bentham in Journal of the Linnean Society, xvii. 189-108, in which a just distinction is indicated between changing a well-established name and giving a new name to a new plant. See American Journal of Science for April, 1870. Mention should also be made of Strickland's Report of a Committee on Nomenclature to the British Association in 1842, of Agassiz's classical preface on the nomenclature of genera in his Nomenclator Zoologicus, and of Dall's thorough and well-digested Report of the Committee on Zoological Nomen- clature to the American Association for the Advancement of Science, 1877, — these dealing primarily with zoology. 348 PHYTOGKAPHY. that ma}- create error or ambiguity, or throw confusion into science. Next in importance is the avoidance of any useless introduction of new names. Other considerations, such as absolute grammatical correctness, regularity or euphony of names, a more or less prevailing custom, respect for persons, &C., notwithstanding their undeniable importance, are relatively accessory.' (Alph. DeC'andolle. 1. c.) 708. The following are universal rules in scientific nomen- clature : - 1. Names must be in Latin or be Latinized. Those from the '.< (sig- nifying likeness) to generic names not of classical origin. Ex. nlulilolihx, lii.rniili s, ttiii-ii/linirlt'S, fnrlisi»ii/i s. 1/1 iiliniiniili'S, lulu liniiti 8, Iminn fortioidl s. Ill Eng- lish, some hvbrids will perpetuate themselves, as for instance terminology, centimetre, millimetre, Iteaurocracy, Xe. 1 Very many, indeed, are adjectives used as substantives, as. Arenaria, Clnrnriii, Sn/ioiiiiriii, fin/mlii us, Triintalix, and even Gloriosa, MirabfllS, &c. Some two-worded generic names anterior to LinnaMis, such as I >i ns Leoms, Vitis lil" n. llnr-iii />,ix/,iris. remain for sections and species, but not for gen- era. When two words are continent into one, they are not objectionable, as Laitrocerasus, Carlemania (commemorating Charles Leman, Carolus Le- manus), &c. NOMENCLATURE. 349 prominent or peculiar character or appearance, from localities, from the names of persons (especially of discoverers), from indigenous or vulgar names, or even from arbitrary combina- tions of letters. Unmeaning names, if not in principle the best, are never misleading. The main requisite is that they should be euphonious, not too long, and that the}' should be adaptable to the Latin tongue. Characteristic names, when possible, are among the best ; such as Saiiguinaria for an herb with red juice, Hcematoxylon for the Logwood tree, Lithospermum for a plant with stony seeds (or seeming seeds) , Myosurus for a plant with gynoecium resembling the tail of a mouse. Names of this sort do not always hold out well ; for Chrysanthemum, so called from its golden yellow blossoms, now has many white-flowered species, Polyyala is wholly destitute of milk, and many species of Con- volvulus do not twine. Neat anagrams are not bad, such as Brown's Tellima for a genus nearly related to Mitella. Personal generic names are wholly proper when dedicated to botanists, especially to the discoverer of the plant, or to other naturalists, or to persons who have furthered botanical investigation or exploration. Ancient names of this kind have been mentioned, also some of those which commemorate the earlier botanists. (702.) At present, almost every devotee of the science is thus commemorated, from Linnaeus and Jussieu downward. In forming such names, the name of the person, cleared of titles and accessory particles (thus Candollea, not DecandoUea) , takes the final -a or -ia and becomes feminine ; and its orthography is preserved as far as possible, making only necessaiy concessions to euphony and to the genius of the Latin language.1 The Linnaean canon forbade the use of the same generic name in botany and zoology, — a rule now impossible to maintain. Perhaps we cannot pre- vent the duplication of phsenogamous names in the lower Cryptogamia. 1 Thus, we may write Lescuria instead of Lesquereuxia, although Michauxia is the form for the genus dedicated to Michaux, however pronounced. The genus dedicated to Strangways is written Stranvcesia (although Strangwaysia might have been tolerable) ; to Andrzeiowsky, Andreoskia ; to Leeuwenhock, Levenhookia (although the elder DeCandolle restored all the vowels), &c. As specimens of overdone simplification, there is Gundelia, named for Gun- delsheimer, and Goodenia, named for Bishop Goodenough, although Gundels- heimera would not in these days be objected to, and Goodenovia is faultless. Yet the names having been so introduced into the science should remain, fixity being of more importance than perfection. Mistaken orthography of the name itself may, however, be set right. Brown's Lechenmtltia is Les- chenaultia, Nuttall's Wisteria (named after Dr. Wistar) is Wistaria. The rule laid down in the code as drawn up by Alphonse DeCandolle is: " When a name is drawn from a modern language, it is to be maintained just as it was made, even in the case of the spelling having been niisunder- 350 PHYTOGKAPHY. 710. The et3*mology of a new genus should always be given. Of the Linmean restrictions, one hold-. \ix. that the names of genera are not to end in -oidts, as many of the older names did. 711. Names of Species are commonly and by preference adjec- tives, agreeing with the name of the genus, and expressive of some character, haliit, mode or place of growth, time of flower- ing, or commemorating the discoverer, first describer, or some one otherwise connected with its history. Thus, in the genus /ii/t/in/rii/its, /i. Imlbosus is named from the bulb-like crown or base of the stem ; R. acris, from the acridity of the juice ; R. scele- ntfiis (the accursed), in reference to the same property ; R. repens, from the creeping habit of the stems; 11. jiusillus, from general insignificance; A'. (Hjuatilis, from its growing in water; R. ni- /V///.N-, from living near eternal snow ; R. Pennsylvanicus, from country or State whence it was first made known to botanists ; R. Bonplandianus, in honor of Bonpland, one of the discoverers ; and so on. More commonly, when a discoverer or investigator of a species is commemorated in the name, this is a substantive, in the genitive, as Ranunculus Nuttallii, i. e. the Ranunculus of Nuttall, instead of R. NuttalliH- ro/fii/i/s si'/tium (of the hedges), Cassia p>n»ili<> (the dwarf) ; more commonly it is a substantive proper name, and this usu- ally an old generic name reduced to that of a species. Ex. llditiniriiliix Flammula, R. Thora, and li. ('t/nilxtlaria; also Lirio- stood by the author, and justly deserving to be criticised." But this is somewhat too absolute, since it is allowed that obvious errors in the con- struction of names of Latin or Creek derivation may be corrected, provided the change does not affect the initial letter or syllable, and that no ancient names are to be disturbed. The clause that forbids changes in the orthography of ancient names, even to make them classical, is a very proper one. The botanical Latin of Tonrnet'ort, Linna-us, Jussieu, and their contemporaries, has by pre- scription rights which botanists are bound to respect. Wherefore fi/ms is the botanical name of the pear-tree, notwithstanding the classical Pirns. So l,i-i -is, as a specific name for a smooth plant (and as distinguished from Irt-is, a light or slight one), is fixed by long botanical use, although only levis is classical ; and it is unnecessary to change Ranunculus acris to R. acer. NOMENCLATURE. 351 dendron Tulipifera, Rhus Toxicodendron, Dictamnus Fraxinella. These proper specific names take a capital initial letter.1 Rarely such a name is in the genitive ; as Heterotheca Chrysopsidis, mean- ing a species of Heterotheca with the aspect of a Chrysopsis. 713. Specific names should be of a single word. Some few are compounded, as purpureo-ccendeum ; and some of ancient origin (once quasi-generic) are of two words. Ex. Panicum Crusgalli, Capsella Bur sa pastor is, Taraxacum Dens leonis. 714. A specific name cannot stand alone. It is nothing except as connected with the genus to which it pertains. A Japonica b}r itself is wholly meaningless. A plant is named by the mention of its generic appellation followed by the specific. 715. Names of Varieties. These are in all particulars like specific names. Many are specific names reduced, to a lower rank. The varietal name is written after the specific, thus : Ranunculus Flammida, var. r»ptans, and R. aquatilis, var. tricho- phylhis. Varieties of low grade need not be named. They may be designated by numbers, or by the small letters of the Greek alphabet, «, p, &c. When the varieties are marked « and j3, the first is supposed to be the type of the species, or both to be equally included in the common character. But when the « is not used, the varieties rank as deviations from the assumed type of the species. Varieties of cultivation, half-breeds or cross-breeds, and the like, should have only vernacular names, at least not Latin ones such as may be confounded with true botanical names. 716. Names of Hybrids are difficult to settle upon any com- plete system. When of unknown or uncertain parentage, they have been named in the manner of species, but distinguished by the sign X prefixed. Ex. X Salix' capreola. Hybrids of known parentage are named by combining the names of the two pa- rents, thus : S. purpureo X dapknoides, or X S. purpureo-daph- noides, for a cross between S. purpurea and S. daphnoides, of which the first supplied the pollen to fertilize the second. The counterpart Irybrid is X S. daphnoideo-purpurea. 1 In respect to the initial of geographical specific names, being adjec- tives, such as Americana, Canadensis, Virginiana, Europcea, Anf/lii-a, usage governs, and this is divided. But the elder DeCandolle, who ruled in all such matters in the preceding generation, always employed the capital in- itial, and two generations of DeCandolle follow the example. Most English authors until recently and some continental ones adopt this usage; and it accords with the genius of the English language, in which we always write European, British, American, &c., with a capital initial. Of late it is a usual practice to write such geographical specific names with a small initial. 352 PHYTOGKAl'HY. 717. The Fixation and Precision of Names. The name of a plant is fixed by publication, and takes its date from the time when it is thus made known to botanists. 718. A genus or other group is published when its name and characters (or the differences between it and all other such groups) are printed in some book, journal, or other adequate vehicle of publication, which is placed on public sale, or in some equivalent way is distributed among or within the reach of botanists. A printed name without characters, and charac- ters without name, do not amount to publication.1 71D. A species is not named unless it has assigned to it both a generic ;iml a specilic name. It is not published until it is made known, by name and characters (or by name along with sufficient information as to its characteristics), in the manner aforesaid. (718.) Adequate distribution, among botanists and public herbaria, by sale or otherwise, of a collector's or distrib- utor's specimens, accompanied by printed or autograph tickets. bearing the date of the sale or distribution (that is, publication by named /•J.'-stccatee in place of printed descriptions), is held to be tantamount to publication.2 720. Characters, references to date and place of publication, and the like, belong to bibliography or particular phytography. not to nomenclature ; but proper identification of names requires that the name of the author and the time and medium of pub- lication should be taken into account. Anterior to the binomial nomenclature, the botanical name of the common tall Buttercup was " Hiimuiculus pratensis erectus acris" according to Bauhin, in his Pinax, p. 171). Under the new nomenclature, which re- duced the specific part of the plant's name to one word, this became Ranunculus acris in Linnaeus, Species Plantarum (ed. 1), p. •">."> 4 ; and a brief character gave its distinctions. In later works it has been more fully described, in some illustrated by figures. The citation of these works arranged in chronological order (or in some order), with reference to volume, page, and in some cases figures, is the bibliography of the plant/' A bot- Names may he communicated, in manuscript or otherwise, by tin- pro- pounder to an author \vho may make them known by publication; but the date of the genii- or other group is that of actual publication. : This does not cover all the conditions of publication, since it docs not specify the characters (and the same may be said of a published figure, with analyses) ; but, on the other hand, it conveys to the competent person receiving the. same all this information and more: so that it should carry the rights of true publication as against any author to whom such names are or should be known. That is, such arc not in the category of " unpub- lished names," which generally ought to be left untouched. 3 For good examples of bibliography, see such detailed works as De- NOMENCLATURE. 353 anist, in referring to this or any other plant, might cite any work which describes it, or none at all. Ranunculus acris by itself, as it happens, would lead to no ambiguity. Not so with many names. For the accurate indication of the species, it is generally needful, or highly convenient, to specify at least the name of the author who first published the adopted appellation. So we write Ranunculus acris, Linn., or L., the abbreviate! 1 name of Linnaeus.1 Here we have the name of the plant, and the bibli- ography reduced to its initial. To this, further citation and other references ma}' be added or not, as the particular case requires. But, so far as citation or reference proceeds, it should simply state the history correctly and clearly. 721. When a species is said to be of Linnaeus or DeCandolle or Bentham, it is simply meant that the adopted name of the plant (consisting of the generic and specific parts together) was first published by this author. Some other author may have named it differently, and even earlier. The earlier name may have been discarded because the specific portion of it was un- tenable, either on account of preoccupation or for other valid reasons. Or the later author ma}^ have differed from the earlier in his views, and have referred the plant to some other genus. As instances of the first, Euphorbia nemoralis, DarL, is a good species, first named by Darlington in his Flora Cestrica. But the name of Euphorbia nemoralis had already been applied to and was the recognized name of a different species of the south of Europe. Whereupon, as the North American species had no other trivial name, a new one had to be given to it ; and it was named E. Darlhigtonii, in honor of the discoverer and first describer. The common Milkweed of Atlantic North America was named by Linnaeus AscJepias Syriaca. As this plant is not indigenous to any part of the Old World, and does not at all inhabit Syria, this trivial name is not merely faulty but false ; so it was changed by Decaisne into A. Cornnti. in commemora- tion of an ante-Linmean botanist who collected it in Canada and gave the first account and figure of it. As an instance of the second, take the pretty little vernal plant Anemone thalictroides, L., meaning an Anemone resembling a Tlialictrum. When it was seen that the essential characters were rather those of Tha- lictrum, the plant was placed in the latter genus. This was first clone in Michaux's Flora ; and so the accepted name is Tlialictrum Candolle's Systema Vegetabilium, and Scrcno Watson's Bibliographical Index to North American Botany, in the Smithsonian Miscellaneous Col- lections. 1 For Abbreviations of Authors' Names, see 385. 354 rilYTOGRAPHY. unemonoides, Michx., meaning an Anemone-like Thalictrum. and Michatix is tin- authority for this name. The names which for any reason are supei>eded become Synonyms* (755.) I'l'l. A later author may circumscribe a species or a genus diH'ereiitly from the originator of the name. To a greater or less extent, this must continually happen in the course of time. Kill ••lilt-inns fnntiiiiinis. Linn.," stands unmoved by the sub- sequent admission of various species (known or unknown to Linmeus) and the linal red net ion of all to one by a thorough monographer. So does Silene Gallica, Linn., although S. quin- guevulnera, Linn., of the same date, is reduced to it. There is no sullieient i-eason for writing Myosutis, Broini. or < ////«y«.v\////<, Brown, because this author restricted the limits of the-e -vuera; nor to write Cilia, Bent/,., because lieiitliain vastly extended 1 Tlie synonymy is an essential part of tin- bibliography or scientific history of a genus or species. But synonymous and admitted names ought to be kept di-tinct. Keeping this principle in view, — also the decisively affirmed doctrine of the founder of our nomenclature, that the specific name is a nullity apart from the generic (so that only the combination of the two makes I he name of the plant, as truly as the constituent halves make the scissors), and bearing in mind the fundamental importance and absolute- ness of the rule that no new names ought to be made where there are tena- ble old ones, — the student need not be misled by the confusing (however specious) innovation countenanced by many /nul.iLiiMs and some botanists, and which has of late years been very fully di.-cu.-scd. The true rule is : " For the indication of the name or names of any group to be accurate and complete, it is necessary to quote the author who first published the name or combination of names in question." (A. DC ) Thus, Leon/ir, //Hilii-fi-on/is, /.inn., fulfils the condition, except where a reference to the work as well as the name of the originator of the name is demanded. Then the citation would continue, "Spec. PI. 312," and might be further ex- tended. In the Flora of Micliaux, this plant was treated as distinct from I.ennticc in genus; and some botanists adopted this view, while others of equal authority did not. Those who adopt Micliaux 's genus name the plant Caulophyllum t/m/irti-oiilix, Mit-luc. Nun some naturalists quote for the species the author who originated the trivial appellation even when transferred to another genus. They would adopt the genus Caulophyllum, yet write : (',iiil(>{>fii//liini tl,l,i/lliini tlinln-tmnl, s (I.iini. sub l.i untie, } Mii-li.i-.," or " Caulophyllum (Michz.) t/Hilii-trai'f/ix, Linn, sub L* nnti<; ," or "('•iti/nti<-t , I. inn.) ilinHctroides, Mi<-li.r." All such endeavors to mix synonymy with nomenclature appear to be faulty in principle and unwieldy in practice. In the most abbreviated form, they state that which is not true: in the others, they impair the sim- plicity and bre\ ity of the binomial nomenclature. It is all but certain that, if the genus ('aulophyllum had been published in the lifetime of Linnaeus, hi1 would not have adopted it. NOMENCLATURE, 355 the comprehension of this genus. Yet in their proper place such changes may be indicated by " pro parte" or " char, muta- tis" " excl. s/J.," and the like, -- useful qualifying statements, but no part of the name. 723. Exactness requires that when a group is changed from a higher to a lower rank, or the opposite, the name of the author who made the change should be quoted.1 He alone is responsible for it. But this rule has only recently been strictly observed. 724. In transferring a species from one genus to another, its specific name must be preserved (with alteration of the gender, if need be), unless there is cogent reason to the contrary. It must necessarily be changed when there is already in that genus a species of the same name ; and then synoirymous names of the transferred species have their claim in order of date. But whatever name is first employed under the accepted genus, being unobjectionable, should hold, even against an older unobjection- able one coming from a wrong genus. This is an application of the stringent rule that no needless names should be created.* 1 Thus, Potentilla Canadensis, L., var. simplex, Torr. Sf Gray, and not of Michaux, for it is the species P. simplex, Michx. Geum, subgen. Stylipus- Torr. Sf Gray, not of Raf., for it is the genus Stylipus of Rafinesque, who neither made the subgenus nor approved it. So, also, for the genus Labur- num we write "Laburnum, Griseb. ; " for even if it exactly corresponded with Cytisus sect. Laburnum of DeCandolle, the latter is not a group of equiva- lent rank. But, as to genera and subgenera, this precision should not be insisted on for times quite anterior to the recognition of such rules and of their need. Spergularia began with Persoon as a subgenus in the year 1805, and this date has been assigned to the genus, although it was taken up as such only in 1819 by Presl and in 1824 by Bartling. 2 Thus, in the case of an older specific name being known, as that of C/ii/o/isis saligna, Don, recognized as Bignonia linearis, Cav., though Don ought to have adopted the latter trivial name, yet as he did not (and the rule was not then really in force as now), there was no need for the introduction of a third name, Chilopsis linearis, DC. " So, again, an Indian Grass was first named and described by "Willdenow as Coix anmdinacea, then named by Roxburgh as Coix barbata, and entered in Sprengel's Systema with Willdenovv's character as Coix Kcenigii. All these names were defective as referring to a wrong genus. Brown corrected the error by creating the new genus Chionachtic, and selected Roxburgh's specific name as the one most generally known and the least liable to misinterpretation , and Brown's Ckionachne barbata is therefore the first correct name ; for which Thwaites afterwards substituted Chionachne Kctnigii, an entirely new and useless name, which falls by the law of priority. It should be well borne in mind that every new name coined for an old plant, without affording any aid to science, is only an additional impediment." Bentham (Notes on Euphorbi- acese, in Jour. Linn. Society, xvii. 197, 198, November, 1878). The following 356 PHYTOGKAPHY. 725. Names of Snbgenera or of other sections of genera are like those of genera ; indeed very many of them, and the most lilting, are old generic names which ha\e been comprehended in the genus by reduction. Unlike genera and higher groups, however, sections, when of (i reek derivation, may properly take the termination in -aides,1 and the typical section may l>cur the name of the genus with the prefix tin.- Sections need not be named at all, and only those of comparatively high rank should is a farther extract i'rmu the same protest against the practice "of creating anew name in order to combine an old specific with a new generic one: " " In Ferns, the wanton multiplication of ill-defined or undefinable genera, according to the varied fancies of special botanists, has had the effect of placing the same species successively in several, sometimes seven or eight, different genera ; and it is proposed to maintain for the specific appellation tlii' right of priority, not only in the genus alone in which it is placed, but in the whole of the genera to which, rightly or wrongly, it has been referred. This has been carried to such an extent as to give to the spccilic name a general substantive aspect, as if the generic ones were mere adjuncts, — a serious encroachment on the beautiful simplicity of the Linnajan nomen- clature; and it is to be feared that there is a tendency in that direction in pha'iiogauiic botany. When a botanist dismembers an old genus, the rule requires that he .should strictly preserve the old specific names in his new genera; and, when he has wantonly and knowingly neglected this rule, it may be right to correct him. But where a botanist has established what he believes to be a new species, and has therefore given it a new name, the changing of this name after it has got into general circulation, because it has been discovered that some other botanist had previously published it in a wrong genus, is only adding a synonym without any advantage what- ever, and is not even restoring an old name; for the specific adjective is not of itself the name of a plant. ... A generic name is sufficiently in- dicated by one substantive; for no two genera in the vegetable kingdom are allowed to have the same name ; but for a species the combination of substantive and adjective is absolutely necessary, the two-worded specific name is one and indivisible; and combining the substantive of one with the adjective of another is not preserving either of them, but creates an abso- lutely new name, which ought not to stand unless the previous ones were vicious in themselves, or preoccupied, or referred to a wrong itl<:t or -O/M.-/.S, as Axt< rn!
  • (p, Filices. and even Aroideee and Ficoidece, will retain these appellations ; but no new ones of the kind will be made. 7:!1. Also, names formed from genera which do not well take the termination in -accce may be allowed as orders to retain their natural form in -inece, -idea, -ariece, and the like. Ex. Tamaris- clnece, Salicinece, Scrophularineee, Berberidece, Lentibulariea. We may prefer for the sake of uniformity to write Salicaceez, Berberi- dacece, Lentibulariacece , and Scrophulariacece (as we should write Violacece} , but this form cannot be insisted on. On the other hand, a termination in -acece has been allowed in the names of certain tribes to avoid excessive iteration of vowels. Thus, for the tribe of which Vernonia is the leading genus, authors write Vernoniacece, to avoid Vernoniece, which ends with four vowels. Spinea and Staphylea are the types of tribes, for which the names, it' they followed the rule, would be Spirceece and Staphyleea, ending one in five the other in four consecutive vowels. Some avoid this l>v writing StaphyleacecB and /Spirceacece. Others write Staphyhee, but this is only the plural of the generic name. I'.Vl. A few orders or other groups took their names long ago from superseded generic names. Ex. Caryophyttacets or < '»/-//<>ji/i////<'(t!, Onagracece or Onagrarieee, and Lentibvlariece. 733. Names of Cohorts are distinguished by the termination in -ales. This was proposed by Lindley, and is adopted by Bentham and Hooker in the Genera Plantarum. Ex. h'foHOcnf>//i't/i>»es. The names of the two great series or sub-kingdoms, following the analogy of the Linn.-ean classes, end in -fa, and are P/uEnogamia, or Pltancroyuiiini, and Oryptogamia. GLOSSOLOGY. 359 SECTION II. GLOSSOLOGY OR TERMINOLOGY.1 735. This is nomenclature as applied to organs or parts and their modifications. The actual botanical terminology owes its excellence in the first place to Linnaeus, and then to DeCandollc. The Theorie Elementaire of A. P. DeCandolle (the first edition of which was published in 1813) is still classical authority, and until recently has received few additions as regards terms need fill in pluenogamous botany. 736. The fundamental rule is that each organ or part shall have a substantive name, and that modifications of organs shall be designated by adjective terms. These names or terms should be as precise as possible : each object ought to be known b}- only one name, yet synonyms are unavoidable ; and no term ought to be used with two different meanings. The word flower, for instance, must not be used for a cluster of flowers, however it may imitate the appearance of one, nor for the corolla or other portions of a flower. Still, some terms have to be used in two or more senses, to be determined only by the connection, or else as having both a special and a more general meaning. Leaf (fo- lium) is a notable instance. A bract, to go no farther, is a sort of leaf; and the imperfect stamens of a Catalpa-flower and Pentstemon are stamens, although likewise called staminodia : these are liable to be called sometimes by one, sometimes by the other name. But, however frequent such ambiguities may be in morphological treatment, they are usually avoidable in descriptive botany, in which terms are held to their more special or partic- ular sense. Yet no rule can absolute!}' determine whether leaf or bract, bract or bractlet, is the proper term in many cases. Moreover, substantive names must also be applied to certain mere modifications of the same organ. In the same family, a simple carpel, differently modified in fruiting, is an akene in a Ranunculus, a follicle in Aquilegia, a berry in Hydrastis and Ac-tea ; while in another family an additional line of dehiscence makes it a legume. Moreover, in this latter family it is called a legume when it is not dehiscent at all, and even when it becomes a drupe ! Arbitrary rules cannot absolutely fix technical any. more than ordinary language. 737. Experience and judgment must determine what modifi- cations of organs should be regarded as a kind, and bear sub- 1 Although the former is the better name, the latter is well established in use as an English word, and perhaps it need not be objected to, inasmuch as the Latin terminus comes from the Greek reppa, of the same meaning. 360 PHYTOGRAPHY. stantive instead of merehy adjective names. But the former should not be unnecessarily multiplied. 738. The classical language of scientific botan}' being Latin, all the organs of plants and their principal diversities are desig- nated by a Latin or Latinized name. .Modern languages have also their own names and terms. (1 really to its advantage, English botanical terminology has adopted and incorporated terms from the Latin and ( ireek, with slight changes, not obscur- ing the identity, thus securing all their precision, and rendering the simple botanical Latin of descriptions of eas}* acquisition to Hie English student. 739. In a text-book like this, the principal names and terms applied to organs and their leading modifications, as also those which relate to their action (physiological terms), or to our study of them (didactic terms, such as phytography. phyllotaxy. glos- sology), art- defined and illustrated in course. There remain the more numerous and varied characteristic terms, chiefly adjec- tives, applicable to more than one or to all organs, and which compose the- greater part of glossology. These, which DeC'andolle arranged systematically with much elaboration, may best be reached by a glossary or dictionary, such as that at the end of this volume, which comprises the substantive terms likewise. 740. From characteristic adjective terms are derived the greater number of specific names of plants ; of which, therefore, the glossary may elucidate the meaning. 711. Capable as the existing system is, it cannot in single words define all observed forms and grades, nor well avoid various ambiguities of meaning. Some defects of the first kind are remedied by combining with a hyphen two congruous terms to denote an intermediate state. Ex. ovato-lanceolutus, or ovate- Ifii/i-i'n/ttie, for an outline between the two. Also a term may be qualified by the prefix sub, in the sense of somewhat, as in sub- mt a minx, subcordcUus (somewhat round or slightly heart-shaped), or diminutives (such as integriusculus) , or superlatives (integer- run us) or other strengthened forms (such as peranyustus} may be employed. Among terms of more than one form of meaning are such as cnli/rlmis. which may mean, according to the context, pertaining to the calyx, or of the appearance of calyx : ci/mntns may mean in cymes, or bearing cymes, or in the manner of a cyme : and /m/i'm-rns may mean provided or beset with chaff, or resembling chaff in texture. Often the form of the word should distinguish the sense: as fnliafus, furnished with leaves, foliosus, with abundance of leaves, while fiK may mean either bear- ing leaves, or properly of leaf-like texture1 or appearance. DESCRIPTION. 361 742. Absence of an organ or quality may be expressed by means of a prefix with privative signification, as indehiscent, not dehiscent, exanradate, destitute of a ring, apetalous, without petals. But the Greek privative « should not be prefixed to Latin words, nor the Latin sub to terms taken from the Greek. 743. When the Latin preposition ob is prefixed to an adjective term, it means obversely ; thus obcordatus is cordate inversed, that is, the broader end with its notch at the apex (instead of the base) of the leaf or other plane organ. SECTION III. DESCRIPTION. 744. Under this head may be conveniently comprised all that relates to the form of the exposition, in botanical terms, of the differences by which the species and groups of plants are distin- guished and recorded, the structure exemplified, and the history or bibliography indicated in systematic works or writings. Lin- nanis, in the Philosophia Botanica, treated these topics under the head of " Adumbrationes." 745. Descriptions may be full and general, comprising an account of all that is known of the structure and conformation of a plant or group, or rather all that is deemed worth recording, or they may be restricted to what is thought most important. In the former, the description is independent of all relative knowl- edge, or takes no notice of relationship to other plants or groups. The latter intends to portray the species or group in its relations to others, and to indicate the differences sole!}'. Exhaustive descriptions of the former kind are seldom drawn up, but partial or supplementary ones are common. Descriptions of the latter kind, when reduced to what is essential or differential, are termed Characters, or the Character, of the group so described. There are all gradations in practice between characters and descrip- tions ; but the distinction should be maintained. 746. Characters are specific, generic, ordinal, &c. They are the differentia, or marks which distinguish a group from any related group of the same rank with which it may properly be compared. According to the occasion and purpose, they ma}' specify only the fewest particulars which will serve as a diag- nosis, or they may be extended to all the known constant differ- ences between two or more related species, genera, orders, &C.1 1 The former would answer to what have been termed differential char- acters, the latter to f.wiitictl characters. Linnaeus divided (generic) characters into factitious, essential, and natural; by the former denoting any difference which may effectively distinguish between any two groups brought arti- 362 PHYTOGRAPHY- "What is now termed the specific character was the specific name with Linnaeus and his predecessor.- : what we call the specific, Linnaeus called the trivial name. (7o:;.) 717. Subordination of characters and the avoidance of vain repetitions require that as far as po»ibh regard being had to the form of the \vork--the ordinal character should contain only what is needful to circumscribe it, and to exhibit clearly its morphology; that the characters of tribes or other divisions should not reassert any portion of the ordinal character, nor tin1 generic character that of the superior groups : and so of tin- sections and subdivisions of all grades down to the species. Equally from the specilic character should be excluded every thing which Id-longs to the generic, or is common to its rela- tives generally, or has been already specified in the section or its subdivisions. So, likewise, of the varieties under the spe- cies. This can be done only by so arranging the species as best to exhibit their relationships, that is, by bringing together or into proximity those of greatest resemblance in all respects, or in the more important respects. AVhat these are, and how a just subordination of characters is to be apprehended, cannot be taught by rules, but must be learned by experience and from the critical study of the classical botanical works. No one is competent to describe new plants without such study, and without a clear conception of the position which a supposed new species should occupy in its genus, or a genus in its order. 748. Characters of orders, genera, and of all intermediate groups, are drawn almost without exception from the organs of fructification. In the description, these parts are mostly taken in order, beginning with the calyx and ending with the ovary, the fruit, seed, embryo. But, as to the orders, some writer- pre- fer to preface these proper characters with a general sketch of those derived from the vegetation, which, albeit of less syste- matic value generally, are often very characteristic of particular families. Knbiaccn-. for example, are known by their opposite1 entire and simple leaves and intervening stipules, along with a few floral characters; Sarraceiiiaeeie. by tubular or pitcher-like lea\es. along with a certain combination of a few other charac- fifi.-illy together, as they mi-lit be in nn artificial key, and as very inliko lien era often \vcrc in his sexual sy>tnn ; by the second meaning the distinc- tions, the fewer the 1 let ter, \vh icli will separate- a group from its nearest relatives ; by the third, all real marks of difference, /. e. all afforded by the oruans of fructification, which only were taken into account for genera, &cu I'pon the construction of this natural character I.inna-us prided himself. and justly. These are the characters in his Genera Plantarum. CHARACTERS. 363 ters, and so on. Where brevity is aimed at, such external and obvious characters, followed by a few diagnostic marks, may practically take the place of a full enumeration of particulars, many of which maybe common to other orders, though not in the same combination. Generic characters always commence with the calyx or most external of the floral organs and proceed to the ovary, thence to the fruit and seed, and end with subsi- diary (but often no less diagnostic) particulars furnished by the vegetation and mode of growth. 749. Detailed descriptions of species, as distinguished from technical characters, commence with the root, and proceed in order to the stem, leaves and their parts or appendages, inflor- escence, bracts, flowers, calyx, corolla, stamens, with filament, anther, and pollen, the disk, if any, gynoecium and its parts, ovules ; then the fruit, seed, albumen, if an}-, embryo and its parts. But descriptions of this sort in most works and in ordi- nary cases are partial and subsidiary, comprising onl}- certain details supplementary to or in amplification of the character of the species or genus. In condensed works, such description is wholly omitted, or is reduced to a few specifications which do not readily find their way into the character. 750. Specific characters usually follow the same order of enumeration, from root to seed, so far as the several organs are mentioned ; and in Latin the phrases are expressed in the abla- tive case. But these particulars are often very conveniently- prefaced b}- statements applying to the whole plant rather than to any one organ ; and these are given in the nominative, and agree with the name in gender.1 751. Linnanis required that neither the essential character of a genus, nor a specific character (his nomen speci-ficum), should exceed twelve words. Latin characters take fewer words than English. But this arbitrary rule is wholly out of date. Yet such characters should be brief and diagnostic : otherwise, their advantage is lost, and the distinction between them and descrip- tions disappears. In monographs and floras, the desirable brevit}', or such as the case admits, is secured b}- proper group- ing under a subordination of sections, subsections, and other subdivisions.2 1 Ex. "NEPETA CATAKIA : erccta, data, cano-pubescens ; foliis petio- Jatis," etc. In English, these adjectives without any substantive expressed will be seen to belong, as here, to " plant " or " herb " understood. - In the Synoptical Flora of North America, such a system of successive divisions is thoroughly carried out. And, if the specific characters are by no means short, it is mostly because nearly all separate descriptive matter is 364 752. Punctuation. In proper descriptions, and in characters of genera ami <>f higher groups, the account of each organ forms a separate sentence : and in Latin the terms are in the nomina- tive case, except subsidiary portions, which are often thrown into the ablative. Kxcepting the latter part, the adjective terms are separated b}' commas. A specific character is always in one sentence. In Latin, its clauses are mainly in the ablative ; and much diversity prevails as to the punctuation.1 Suhgvneric and other sectional characters are commonly framed like those of dispensed with: consequently various particulars are added to the char- acter which do not strictly belong to it. In Bentham's great Flora Austra- liensis, also in English, specific characters are replaced l>y a characteristic synopsis at the head of each genus ; and a terse description under each species completes the account. Moreover, Bentham, in recent works, Midi as his revision of the Genus Cassia, also that of the JMirnosea1, which have Latin characters, writes these in the nominative case and each member in a separate sentence, in the descriptive form, abandoning the long-used abla- tive form. 1 Linnaeus employed only the comma in the specific character, along with a subsidiary use of the colon in a manner very unlike its ordinary use in punctuation, making it a point of less value than the comma. Thus, " CHENOPODIUM ALHTM foliis rhomboideo-triangularibus erosis postice intcgris : summis oblongis, racemis ercctis." Spec. I'l cd. '2. 019. Here, while- the two main members of the sentence arc separated by a comma, a subsidiary portion of the first member, relating to the uppermost leaves, is separated by a colon. Limucus employed the colon in the same way in generic characters. This anomalous usage is now abandoned. But most authors have followed the Limiu?an pattern in distinguishing the prin- cipal members by commas only, so that these become the only points in the specific character, however complicated that may be. Thus, " UVM \< i i.i:s ACKIS (Linn. Spec. 77!>) foliis pubescentibus subglabrisve palmato-partitis, lobis inciso-dentatis acutis, summis linearibus, caule erecto pluriHoro subpubescente, peduneulis teretibus, calyce subvilloso, carpellis mucrone subcrecto terminatis " DeCandolle, Prodromus, i. 36. This is the punctuation throughout the Prodromus and in most contem- porary systematic works. Its imperfection is shown in the above-cited speci- men. The primary members of the sentence, which characteri/e the leaves, stem, peduncles, calyx, and carpels, are distinguished by the same grade of punctuation which serves for the parts of the first member, viz. the lobes of the leaves, and for a still subordinate portion, viz. the form of the upper- most lobes. This want of subordination is to be remedied by the use of semicolons between the principal members, and of the commas only fi>r the secondary ones, — a punctuation now not uncommon, and which is adopted in the recent tirst volume of the Monographic Phanerogamanun of the I)e- Candolles, which supplements the Prodromus. The portion of that volume contributed by Dr. .Masters better exemplifies this than does the rest of the volume. For the latter sacrifices the advantage of the change by the inser- tion of commas between each adjective of a continuous ablative phrase (as, " Srnilax laurifolia ; limbis foliorum oblongis vel ovato-oblongis, coriaceis, 3-6-nerviis, subtus pallidioribus," etc.), where they are generally deemed PUNCTUATION, SYNONYMY. 365 genera. Or the members may be united in one sentence, but in that case the principal ones are best separated by colons. 753. Should a point intervene between the specific name and that of the author cited? The practice varies. But, if the name is Latin, the comma is superfluous ; for the abbreviated name of the author is supposed to be in the genitive, and to read thus : Ranunculus repens Linncei. Still, since when the author's name is cited in full it is never written in the genitive, and since in English the comma is normally required, it seems on the whole proper to insert it. 754. In citations, the classical practice is to separate the refer- ences from each other and from the name by periods ; thus, " Anemone cylindrica, Gra}*, Ann. N. Y. Lye. 3. 221. Torr. & Gray, Fl. 1. 113," l&c. It is becoming equally customary to separate the several citations by semicolons, thus bringing all the references under one name into one sentence. The bibliography of a species or group of species which a describer or other author has to refer to (with more or less fulness, according to the form of his work) is to be sought parti}- under the admitted name, and partly in the 755. Synonymy. This includes all other than the admitted names. Ex. Hydropeltis of Michaux is a synonym of Brasenia of Schreber, the latter being the earlier published name. Nectris of Schreber is a synonym of Cabomba of Aublet, the latter hav- ing priority. Thalictrum anemonoides of Michaux has for syno- nyms Anemone thalictroides of Linnaeus and of many subsequent authors who followed him in referring this ambiguous plant to Anemone (721) ; and also Syndesmon thalictroides of Hoffmannsegg and Anemonella thalictroides of Spach, who proposed to consider superfluous. The preferable punctuation of the character above-quoted from the Prodromus would be RANUNCULUS ACRIS (Linn.) : foliis pubescentibus subglabrisve palmato- partitis, lobis inciso-dentatis acutis, summis linearibus ; caule erecto pluri- floro subpubescente ; pedunculis teretibus ; calyce subvilloso ; carpellis mucrone suberecto terminatis. The advantages of this style of punctuation will more and more appear, when applied to less simple cases. Commas between the ablative adjectives are superfluous and confusing. In English characters, commas are required between the adjectives which follow the noun. Rightly to express the subordination of characters, the plan adopted in the Synoptical Flora of Nortli America is recommended ; that is, with colons separating the principal members, semicolons for subordinate and dependent ones, and commas between the adjectives of the same noun. 1 See Watson's Bibliographical Index to North American Botany (where this style is adopted) for a general model for the arrangement of synonymy and citations. 366 PHYTOGKAI'HY. it an intermediate genus between Thalictrum and Anemone. In systematic works, the specific character inn Mediately follows the name, and generally forms a part of the same sentence: and is followed lirst liy citations of authors who have adopted the name. and then by the synonymy, or as much of it as the plan of the \\ork calls for. The synoin nn.iis names and the references under them should be cited in the order of their publication. But, to economize space, all the authorities for the same name are brought together into one sentence, and arranged according to their dale. Also, where the synonymy is not elaborately displayed, the various synonyms of the same generic name are usually placed in consecutive order. 7 .')i;. Iconography. The leading and most essential citation is that of the author by whom and the work in which a plant is named and described, and also the work in which it is best char- acterized. Among the characterizations, published ligures hold a prominent place. The citation of these is an important part of the synonymy. The best botanical plates are those which give detailed analyses of the parts of the flower, fruit, and seed, displaying their structure. 7.»7. Habitat and Station are recorded in a sentence or para- graph following the name, character, and synonynn of a species. The liiililintlnn is the place, district, or region at or within which the plant is known to be indigenous, or to grow spontaneously. The complete habitat is the geographical range. The station is the situation it all'ects, whether in water, in marshes, on shores, on hills or mountains, in forests, on open plains, &c. 7f>N. Discoverer, \c. To the' habitat and station of newlv • li-covered. rare, or local plants should he appended the name <>!' the discoverer or the collectors by whom the species has lie- come known to science, at least when the plant is first published. Dale of discovery should also then be indicated. 7.".'.). Time of IJIossoniing should be recorded, either the month or the season, to which may be added that of the maturity of the fruit. When the month or season is mentioned without farther explanation, (lowering-time is intended. In a flora, this mav sometimes be indicated under the genus for all the specie-. In the flora of an extensive region, and in respect to species of considerable range in latitude or longitude, the time of (lowering differs so widely at tin- extremes of the geographical range that it cannot well be specified except in general terms, as spring, xii/tn/ii-r, ant n in ii. \c. 7i;o. Ktymology of Names. When a new generic name is pub- lished, its origin and meaning should always be given, if the ACCENTUATION, ABBREVIATIONS. 367 nature of the publication will allow it. So likewise of species, except where the source or signification of the name is mani- fest. This is commonly the case as respects most characteristic specific names, and also those drawn from station, habitat, and the like. 761. Accentuation of Names. The pronunciation of botanical names is settled by the rules of Latin prosody. All that is usually attempted in those botanical works which take this into account is to mark the S3'llable upon which the principal accent falls. This in words of two syllables is always the lirst ; in words of three or more S3'Hables, either the penult (the last sylla- ble but one) or the antepenult (next preceding syllable) . When the penult is a long syllable, it takes the accent ; when short, this recedes to the antepenult. The accentuation may accordingly be sufficiently indicated by marking the quantity of the penult, either long as in Erica, or short as in Arbutus and Gladiolus. Or else the accent may be marked by a proper sign, as Erica, Arbutus, Gladiolus. An endeavor has been made to represent the longer sound of the vowel by the grave accent-mark, as Erica, and the ohort by the acute, as Gladiolus. But this plan is encumbered with practical difficulties. 762. Abbreviations are required, both of the name of the au- thor, when of more than one or two syllables, and of the titles of the works cited. There are also the customary abbreviations in the citation of volume, page, plate, &c., in which there is nothing peculiar to botany. 763. The simple rule for the abbreviation of an author's name is to abridge it of all but the first syllable and the first letter of the following one (as Lam. for Lamarck, Hook, for Hooker), or the first two letters following the vowel when both are consonants (as Linn, for Linnaeus, Juss. for Jussieu, Rich, for Richard). Sometimes more of the name must be given, in order to distin- guish those beginning with the same syllable. So we write M'n-h.f. to prevent confusion of the name Michaux with that of Micheli, which, being the earlier, claims the abbreviation Mich., and Bertol. to distinguish Bertoloni from Bertero. Sometimes a much-used name of one syllable is abbreviated, as Br. (or R. Br.} for Robert Brown. Initials or abbreviations of the bap- tismal name are needed to distinguish botanists of the same name ; as P. Browne in distinction from Robert Brown, Ach. Rich., Adr. Juss., Alph. DC., to distinguish the younger from the older Richard, Jussieu, and DeCandolle. Or, where father and son, the abbreviation for the latter may be Jitss. fL, Hook, fil., or Hook. /., &c. Certain, but very few, well-known and eminent 368 PHYTOGRAPHY. names are abbreviated to a sign ; as L. for Linnaeus, DC. for DeCandolle, HBK. for ilumboldt, Bonpland, and Kunth, the latter too long after ordinary abbreviation.1 Care should be taken to ailix the period by which abbreviations may be dis- tinguished from full names, siieli as Don, Ker, Blytt. 7U 1. Alibreviations of titles of works follow the same rules as those <>f names, or at least are in no wise peculiar in botany. 7(;."». Abbreviations of the names of organs follow the same rule: dil. for calyx, C<»-. for corolla, Slam, for stamen or stamina, Pi'st. for pistillum or pistil, Fr. for fructus or fruit, Per. for pericarpium or pericarp, Sem. for semen or seed, are the most common. Hab. for habitat or geographical station. Herb. for herbarium, Gen. for genus, Sp. or Spec, for species, Var. for variety, and the like, every one will understand. But some abbreviations which are common in botanical writings, at IcaM those in Latin, may need explanation to the elementary student. A list of abbreviations is appended. See p. 390. 766. Signs. Under this head might be ranked such abbrevia- tions as v. v. for vidi vivam, v. s. for vidi siccam, to note that the writer has seen the plant, either alive or in a dried specimen ; or, more particularly, /•. s. s., when it is a spontaneous specimen that has been examined in a dried state, and v. s. c., when it was a cultivated specimen ; v. v. c., when the living plant was seen in a garden only, and /•. v. s., when the spontaneously growing plant was seen alive. There arc also proper signs, of which the most common are those which indicate the sexes of blossoms, the duration of a plant, and the like. Also the interrogation point (?) used to express doubt ; the exclamation point (!) to indicate the certainty that is given by the actual sight of an authentic original specimen. Seep. 3!>1. 7(')7. The marks r.sed to indicate- the subordination of sections under a genus, or in the synoptical arrangement of genera, and the like, are not settled by any fixed rule. An approved ar- rangement is to employ the following marks in the given order, §*•»-++=. The first one. for sections of the highest order, takes numerals after the sign. Ex. § 1, and so on. When 1 A- Ahiii DeCandolle remarks, the proper abbreviation of the name he lic.-irs is C,n,il. Rut the form DC. was very early adopted by the first of the illustrious name, and lias he'-n continued for almost throe quarters of a century. Alphonse DeCandolle would prefer to write it D.C., but has not adapted that mode, nor should we; for DC. and 1IBK. arc convenient ab- breviations reduced to si<_riis. Rut such forms should not he increased. For ordinary name* they would he unintelli'jihle. Names which are not too loiiir, ar.d of which an abbreviation by tbe ordi- nary rule is insufficient, such as Decaisne, should rather be written in full. SIGNS, ETC. 369 such sections are followed by a substantive name, they are equivalent to subgenera.1 Ex. Phacelia, Juss., § 1. Euphacelia, i. e. the true or t}"pical Phacelia ; § 2. Cusmanthus, Gray, &c. Sections next in rank to these are marked with asterisks, * for the first, * * for the second, * * * for the third one of the same rank. Divisions of these have the -i— prefixed ; and so on in the same way. Still farther subdivisions may be marked by the small letters of the alphabet consecutively, a, b, c. When capital letters are used for division marks, it is mostly for those of a high grade. 768. Floras, Monographs, &c. A systematic work describing' in proper order the plants of a country or district is generally called a Flora. A Flora of a small district takes the diminutive name of a Florula. A universal work of the kind when it ex- tends to the species is a System, Systema VegetabiUum or Systema Regni Vegetabilis. The latest completed Systema VegetabiUum is that of Sprengel (1825-1828), in five octavo volumes, on a very condensed plan. A compendious Flora or Systema is often termed a Prodromus, literally meaning a forerunner or preliminary work. But, as even this is more than most bot- anists are able to complete, the name of Prodromus is now applied to works which are not intended to precede fuller ones b}- the same author. The principal work of this kind is the • Pro- dromus Syst. Nat. Regni Vegetabilis, commenced by DeCandolle in the 3~ear 1824, continued by his son Alphonse DeCandolle (aided by various botanists) to its close in 1873, down to vol. xvii., or essentially twent}- very compact octavo volumes, these carrying the work only through the great class of Dieotyledones. But the publication of the monocotyledonous orders has com- menced in a series of Monographs (Monographic Phanerogam- arum'). A Monograph is a systematic account of all the species of a genus, order, or other detached group. 769. Specimens of botanical characters and descriptions, cita- tions, &c., illustrating this chapter, might be given here. But, for those in Latin, the classical works of DeCandolle and others, and for the genera those of Jussicu, Endlicher, Bentham and Hooker, may be taken as models. In English, those of the latter authors, and in the United States the better-known writings of the present author, especially the later ones, may be referred to. 1 DeCandolle in the Prodromus employed the word Sect. (Sectio) for what answers to subgenus or at least to the highest grade of sections ; then § 1, § 2, &c., for the next grade below subgenus ; and then the asterisk, and other marks. 24 370 PHYTOGRAPHY. SECTION IV. SPECIMENS ; DIRECTIONS FOR THEIR EXAMINA- TION, PRESERVATION, &c. 770. Implements. Those necessary for the examination of })h;vnogamous plants. Ferns, and the like, are a simple pocket lens, a simple dissecting microscope ; also a sharp thin-bladed knife and some needles of various fineness, mounted in han- dles, for dissection. 771. For a single hand lens, one magnifying only from four to six diameters is the most useful. A doublet, or a parabolic lens of Tolles, of about an inch focus, is better, but much more expen- sive. The simple stage-microscope for dissection need have only two lenses (doubl'ts or otherwise) with large field and good definition, one of an inch and the other of about half inch focal distance: and a glass stage of at least an inch and a half in diameter. A compound microscope is useful for all minute investigation, and is essential in the study of vegetable anatomy and of all lower cryptogamic botany. 772. For making thin slices, a razor is the best knife ; for dis- section on the stage of the simple microscope, beside needles, small scalpels or some of the cutting instruments used by ocu- lists are very convenient. But an expert hand is able to do almost every thing with a common knife or scalpel and a pair of mounted needles. Slender forceps are almost indispensable : those made for the use of dentists are the best. 773. Analysis. In the examination of an unknown plant with a view to its determination, its whole structure should be made out, so far as the materials allow, before a step is taken to ascertain its name and place in the S3-stem. In respect to the stem, its duration and consistence and its internal structure, whether exogenous or endogenous, are to be noted. As to the foliage, the venation and the phyllotaxy. also the presence or absence of stipules, are most important. The antkotaxy or inflorescence is to be examined and referred to its proper type. In the llower. the numerical plan and symmetry, its ground-plan and the nature of the deviations from the general or the- family type, are to be considered; also the aestivation or arrangement of the parts in the bud. the character and extent of coalescence and adnation ; the manner in which the anther is borne upon the filament, and its place and mode of dehiscenee. &c. Note also whether, when the blossom is hermaphrodite, the anthers and the stigmas mature at the same or at different periods. The placen- tation and the character and position of the ovules should be SPECIMENS. 371 determined. Two sections of the flower should be made : one of them vertical and directly through the centre, in the manner of Fig. 33G-341, — this will display the adnation, insertion, &c., of all the parts ; the other transverse and through the middle of the ovary, also above the ovary when this is inferior, and if pos- sible in the unopened but full-grown flower-bud ; this, among other things, will bring to view the aestivation. (Fig. 351, 3!).s, &c.) Not rarely fruit and seeds are to be had at the same time, or upon the same specimen, and these are equally to be investi- gated. In fresh seeds, even those of minute size, the embiyo ma}' almost alwaj's be extracted or brought to view under the microscope, either by tearing away the seed-coat with needles or by sections with a keen knife. When hard and dry, they have, only to be soaked or slightly boiled. 774. Diagrams and also sketches of the parts should be made, such as those referred to in the foregoing paragraph. Such diagrams can be drawn by any one with a little practice ; and the}" may be made to express the whole floral structure, even to the coalescence and adnation.1 But in the process of determina- tion the student should beware of trusting wholly to his diagrams and sketches without direct verification. 775. Dried specimens, when well prepared and in sufficient abundance, in the hands of a skilled botanist are in most cases but little inferior to fresh ones. When needed, flowers, or clus- ters of blossoms, or fruits may be detached and prepared for examination and dissection by somewhat prolonged soaking in warm water or by a short immersion in boiling water. This re- stores flower-buds and small flowers and fruits, or their parts, to a condition not essential!}' unlike the living state. Consequently, the Herbarium or Hortus siccus of the botanist is to him more essential than the botanical garden, important as that ma}* be. 776. Herborizing.2 The collector's outfit will essentially con- sist of a Vasculum or botanical box, a Portfolio, a Trowel, a pocket Lens, and a small but stoutly covered Note-book. Some use a1 portfolio only, others the botanical box ; but on a long excursion it is well to have both. The former is preferable in most cases, except when specimens are collected for the immediate use of a 1 See Eit-liler's Bluthendiagramme (Leipsic, 1872, 1878), an admirable work, which may serve as a model. 2 These articles, from paragraph 776 to 802 inclusive, were obligingly pre- pared, at the author's request, by LYMAN H. HOYSRADT, of Pine Plains, New York. They form an abstract or a new edition of a series of notes on the subject which were published in the Bulletin of the Torrey Botanical Club, in the year 1878. 372 PHYTOGRAPHY. class. When well stocked with paper, it is of almost unlimited capacity ; and most plants of delicate texture (as many of the smaller aquatics, and those with fugacious or delicate corollas) need to be consigned directly to the paper in which they are to be pressed, and to lie kept meanwhile under some pressure. 777. The Vasculmn is very useful for holding plants that are to be examined fresh, and for thick roots, large fruits. «fcc. It is made of tin, and should lie of oval-cylindrical shape, about 17 inches long and 1 by G inches wide, and provided with a Unlit strap to throw over the shoulder. The lid opens nearly the whole length of one of the flat sides (15 In" 4.\ inches, with one fourth inch lap), is made to fit as close as possible, and fastens by a simple spring catch. When no portfolio is used, a larger box ma}- be required. Plants may be kept fresh in such a box for many days. Fora several-days excursion, when it is desirable to bring home a large number of fresh plants, a tin chest, made some- what after the pattern of an old-fashioned trunk, will be found \ery convenient. It should be about 21 inches long. 10 inches wide, and 10 inches high to the top of the convex and hinged lid, which forms the whole top, and to which a handle is fitted. 778. A good form of Portfolio is made of two pieces of binder's board covered with enamel cloth, and fastening together with two long straps with buckles. Handles similar to those on a carpet- bag may be attached for carrying. The usual size of portfolio is 18 by 12 inches, but 16£ by Hi inches may be 1 idler, as there would then be little danger of making specimens of too great length for the herbarium. (7«4.) Or tiie back may be of soft leather, an inch or so in width, and a light strap and buckle at the front edge and at each end. The portfolio should contain a good quantity of folded sheets of thin unsized paper, similar to grocer's tea-paper, and of a size only a little smaller than the sides of the portfolio. Very thin manilla paper, or what is so called, is excellent for this purpose, being sullicienlly bibulous and rather strong. 779. The specimens as soon as gathered should be laid neatly in these folded sheets (called specimen sheets), and kept under a moderate pressure in the portfolio. The sheets with the spec- imens tire afterwards transferred to the home press, but the specimens should be left continuously in their sheets through all the changing of driers, until cured. Indeed, the specimens may well remain in the sheets after drying, until wanted for mounting or for exchanging. For fine specimens, the use of this specimen paper is very important. Many plants are so extremely delicate and sensitive that they will not bear the least handling without HERBORIZATION. 373 curling and shrivelling, unless thus enclosed : also without these sheets much time is lost in transferring small specimens one by one from one driei to another in the drying process. 780. For digging up roots, bulbs, &c., a small and. sharp pointed triangular Trowel or stout knife will answer. One of the best "diggers" is made from a large file. Let a black- smith bend the lower half of the blade to a gentle curve, so that the point will be about an inch out of the true line. Grind off the teeth and re-temper the blade. The total length with handle, which is over one third, should be about twelve inches. A leather case may be made for convenience of carriage. The advantages of this strong tool are many.1 781. A Note-book should be carried upon ever}' excursion, in which the station of rare plants, dates, colors, and various par- ticulars which cannot be learned from the specimens, may be recorded on the spot, instead of being left to uncertain memoiy. 782. For most plants, the best time for collecting flowering specimens is in the morning, soon after the dew has disappeared. Vespertine flowers have to be secured earlier, or at nightfall. 783. Care should be taken to have the specimen of the proper size, neither too small nor too large, and to comprise all that is necessary for complete botanical illustration, — flowers, fruit, and leaves, both cauline and radical when possible. Inex- perienced botanists suppose that a small sprig, containing a flower or two with a few leaves, will answer all purposes as a botanical specimen ; but later he comes to know better, and also learns that the flower is only one of the component parts of a specimen, and not always the most important one. In various genera and orders, the fruit is the most distinguishing character- istic, as with the Potamogetons, the Crucifera?, the Umbellifene, and the Cyperaceae. With man}' plants the radicals-leaves, with others the character of the subterranean stem, whether a rootstock, tuber, conn, or bulb, or of the root itself, whether annual, bien- nial, or perennial, becomes important. Consequently, all the organs have their value in an herbarium specimen, and each and all should receive due consideration from the botanist when col- lecting. Specimens may be often secured that exhibit both 1 [There is an English herborizing trowel of excellent quality, with blade six or eight inches long, less than two inches wide, the sides slightly in- curved, the stout shank an inch and a quarter wide, and one sixth of an inch thick : this forms the whole back of the handle, the front of which is a piece of lignum vitae riveted fast to the steel. It is nearly impossible to break it.] 374 PHYTOGRAPHY. flowers and fruit in the same plant, or fruit may be frequently obtained from more advanced plants at the same time. If not, fruit must be collected later, as in case of shrubs and trees, of which generally only a branchlet with flowers, or with flowers and leaves, can be gathered first. But subsequently the fruit and mature leaves, should always be taken, if practicable, from the same individual as the flowers. Of dioecious shrubs or trees, like the Willows, each species should be repiv>t-nted by four pieces: first, the sterile and fertile catkins will have to be obtained, and the respective individuals marked, so that later corresponding twigs with mature leaves, stipules, and fruit may be gathered, and the specimens rightly matched. 7-s-l. A specimen should be so arranged as to be no larger when pressed than can be neatly mounted on the herbarium paper. A slender plant not over three feet in height should generally be preserved entire, root and all. This can be done by bending or partially breaking it at one, two, or three places, and doubling so that the sections will not rest upon each other in drying. If broken twice, it may be neatly arranged in the N form when put in portfolio. Very large herbaceous plants will have to be divided and the parts preserved separately, or, better, take a suitable portion of the upper stem, having leaves, flowers, and fruit, and a convenient part of the lower stem containing radical leaves and with it sufficient root to show whether the plant is an annual, biennial, or perennial. Thick stems, roots, tubers, bulbs, and the like, should be divided or thinned down with a knife, but in such a manner that the original shape can be easily made out. 7.s.~). Carices should be alwa}-s collected when the fruit is full- grown, but not so ripe as to fall away. Ho also should other Cyperacese ; yet it is well to collect also earlier specimens of these in flower. Grasses, on the other hand, should generally be collected soon after they come into blossom. For when mature the spikelets in many species break up and fall away in drying. The culm, leaves, and root of Sedges and Grasses should be preserved, as well as the inflorescence. The root is no less important. Cespitose species should be so collected and preserved as to show the tufted character. The culms of most sedges and grasses act stubbornly when bent for arrangement in portfolio or press, and are not disposed to stay in place. This dilliculty is promptly remedied by (.-rushing with the teeth the angles made by the bending. Or these may be thrust through slits of paper. In drying Sedges and Grasses, very moderate pressure should be employed. HERBORIZATION. 375 786. Some aquatic plants (Algae especially) are so soft and flaccid that, to secure them in their proper shape, they must be placed in clear water and floated out by inserting beneath them the paper on which they are to remain permanently, either the regular mounting paper, or a thinner white paper which when dry can be pasted on the herbarium sheet. If likely to adhere to the sheet or drier above them in the press, a piece of oiled or stearine paper may be laid direct!}' on the specimens to prevent their sticking. Also viscous or glutinous plants which are liable to adhere to the sheets enclosing them may be sprinkled with Lycopodiurn spores, powdered soapstone, or some similar sub- stance. 787. The name of the plant if known, but by all means the locality and date of collection, with any other descriptive re- marks regarded necessary, should be written on a ticket or on the sheet when it is put into the press. Never omit to record the time and place of collection, as a specimen of unknown date and localit}' loses much of its value and interest. 788. Drying Specimens. The chief requisite for good herbji- rium specimens is the extraction of the moisture from the green plant as rapidly as possible under a pressure which obviates brittleness. This is to be affected by placing the thin sheets containing the specimens between layers of bibulous paper, calle.l driers, and applying moderately strong pressure to the pile. For driers nothing can be better than thick blotting paper, except that it is too expensive, and the same ma}- be said of an English drying paper made for the purpose. Equally good driers are made of the thick and felt-like brown paper which, after saturation with coal-tar, is here largely used under the clapboards of wooden houses and under slate-roofing. It is a cheap material, and is to be obtained, cut into sheets of 18 by 12 inches. Or driers may be made of old newspapers or of any soft wrapping paper, cut or folded to the proper size, and stitched (very expeditiously by a sewing-machine), or joined by eyelet paper-fasteners at two corners, in packages of a dozen or more leaves to a drier. It is well to have a large supply of driers and specimen-sheets ready for use. 789. A half dozen or more pieces of thin boards, 18 inches long and 12 inches wide, should be provided. They are used at the top and the bottom of the pile when pressing, and also for dividing it into suitable sections, especially for separating the packages of plants which were put into press at different periods, and dividing up these packages themselves, if too large. For the plants dry better in small sections and with the pressure :;7(J PHYTOGRAPHY. evenly distributed. Hence it is best to have these sections not over live or six inches in thickness, nor should the pile itself be carried too high, never exceeding t\\<> feet. Painted binders' boards ni:iy be used, instead of the common boards, to separate interior divisions. Some botanists use a kind of lattice made of two layers of thin strips or laths, crossing each other. Thi> U -aid to allow free escape of the moisture by evaporation, and so to accelerate drying, as in the case of the wire press. 790. For giving pressure, various ways have been contrived. The Screw-press is convenient and compact, but objectionable, because it does not follow up the pressure as the plants shrink in drying. This objection does not apply to the Lly used, an etllorcseence may soinetinies be Id'l on the surface of the poisoned specimens upon the evapora- tion of the alcohol. Some add to the solution some carbolic acid, at the rate of a fluid ounce to each quart of alcohol. The solution may be applied with a soft brush (one with no metal in its fastening), or by a dropping bottle, or even the specimens may be dipped in the solution placed in a flat porcelain dish. The brush (using a pretty large and soft one) is the most con- venient and ellicient. The moistened specimens should be placed betwc-.cn driers and in shallow piles until the alcohol evaporates. 805. Thoroughly poison all specimens before admitting them to the herbarium. Jt is well to poison all specimens whatever. as soon as they are made or at the close of the botanizing sea- son, as well those intended for exchanges as for the collector's own herbarium. 806. Keep all specimens between sheets of paper, or within folded sheets, not too crowded or overlaid, away from dust, and in a perfectly dry place, so as to avoid mould. When attacked by mould, the corrosive-sublimate solution should be applied. A properlv dried specimen, duly cared for, should be as lasting as the paper which holds it. 807. The Herbarium, called by the earlier botanists ffortus Siccns, is a collection of dried specimens, named and systemat- ically arranged. It is indispensable to the working systematic botanist, and every devotee of botany should possess, or have access to an herbarium containing representatives of the plants of the immediate vicinity or district, if not of the whole country. Or an herbarium may be restricted to a particular family of plants, made the object of special study. A general herbarium should contain specimens representing all the natural orders and as many of their genera and species as possible. sos. The form of the herbarium as to the size of its sheet- is considerably variable. That of Linna-us is of the si/e of fool-cap paper: this would now be universally regarded as much too small. The principal liritish herbaria adopt the size of 16£ by in.1, indies, which is rather too narrow, rarely permitting two specimens of the same species of any considerable size to be placed side by side on the same sheet. In the United States, 16i inches in length by 1 1 -4! in width is adopted ; that is. for the genus-covers, the species-paper being a quarter of an inch nar- rower. THE HERBARIUM. 381 809. The specimens representing each species may either be laid within a doubled sheet, loosely (as in some European her- baria), or fastened in place by narrow slips of gummed paper (which is much better) , or else they may be glued bodily to single sheets of strong and stiff white paper. 810. The former is an excellent plan for a limited collection. It is an advantage that a specimen can be taken up and examined on all sides ; also, that indifferent specimens can at any time be exchanged for better ones. But a large herbarium on this plan becomes cumbrous and inconvenient for ready reference and comparison. 811. The best plan in a large herbarium, and one much to be consulted, is to attach the specimens completely, by any kind of strong and light-colored glue, to single sheets, or rather half sheets. The specimens are thus safe from injury under reason- able handling, and can be turned over and examined with as much facility as a series of maps or engravings. The species- paper should be of writing-paper stock, or of equal firmness, of compact texture, well sized and calendered, and of a weight in size of 16]- by 1H inches of about 18 pounds to the ream of 480 flat sheets. The paper should be furnished square-cut on all sides, in the manner of " flat cap." Stiffness is the great desideratum. 812. In no case should more than one species be knowingly attached to the same sheet. But of very many species there will be room for more than one specimen. And specimens from dif- ferent localities, of different forms, and in various stages of flowering and fructification, are always desirable. The full name of the plant should be written at the lower right-hand corner of the sheet, or a ticket should there be attached by glue or traga- canth paste. Each specimen should have its ticket, similarly attached, or a memorandum upon the sheet, indicating the hab- itat or the special locality, date of collection, name of collector, and any other desirable information which the specimens them- selves do not furnish. When there are loose flowers or fruits, or when any of these have been detached for dissection and micro- scopical investigation, it is well to preserve them, placing them in little paper pockets or envelopes and pasting these upon the sheet close to the specimen to which they pertain. Sketches of parts dissected may be drawn upon the sheet. Notes and mem- oranda received with the specimen or too extended to be entered upon the sheet may be folded, inserted in such envelopes, and made fast to the sheet. Many botanical collections are distrib- uted with printed tickets. These, and all authenticating tickets 382 PHYTOGRAPHY. or notes, should he attached to the sheet near to the specimen they belong to. In view of this, printed and written tickets should be of small size.1 A ticket whk-h exceeds four by two 1 All printing on an herbarium ticket should be in plain type; and fancy borders, uselessly occupying room, should be avoided. If any border is thought needful, it should be of plain lines. It is not desirable to parcel out the space on a ticket with separate' lines and headings for habitat, date of collection, time of flowering or fruiting, name of collector, and the like. Tlir^e particulars may conveniently be entered at the bottom or top of the ticket, as may be convenient, leaving the rest of the space free for the name of the plant, the authority, and perhaps a synonym. Tickets for specimens distributed among other botanists may well have a head-line indicating the source, such as "EX COLL. c. c. PARRY," or "EX HERB." or, in English, " FROM THE HERBARIUM OF" the botanist who communicates the specimen. The following may serve as an example of a simple ticket for the sending out of dried specimens, and of the way in which the ticket may be filled out with the name of the plant, its habitat and station, name of collector and time of collection. EX HERB. A. GRAY. y*-*~ o, 9 For the botanist's own herbarium, it is well to use a blank ticket with a printed heading like the specimen above, but with the " ex " omitted. When a considerable collection is made in any particular botanical explo- ration or excursion, and numerous or several specimens of the same species are gathered, to be distributed among botanists in the way of exchange or otherwise, these are commonly given out under numbers and with a printed heading to a special ticket. The following is an approved form of such a ticket, and of the mode in which it may be filled up in writing by inserting the name of the species, the locality, &c. No. ALPINE FLORA OK THE ROCKY MOUNTAINS. Coll. C. C PARRY. 1872. COLORADO, ft/VYM/W CY £&<-** --f THE HERBARIUM. 383 inches is a nuisance ; and those of an inch and a quarter or an inch and a half in width and three or four inches in length are most commodious. 813. The sheets of all the species of the same genus, when not too numerous, or of a particular section of it, or any conven- ient number, should be consigned to one genus-cover. The best genus-covers are of manilla-rope paper, the " bleached manilla " such as that of which tags are made is the neatest article, but rather more expensive : they are in whole or folded sheets (pref- erably in quarter quires), accurately trimmed at top, bottom, and front edge to the size of 16 i by 11£ inches; that is, the folded sheet as used is a trifle longer and a quarter of an inch wider than the species-sheets it holds. The sheets to be firm enough should weigh H or If ounces each, or from 45 to 52 pounds the ream. The generic name should be written in a bold hand on the lower left-hand corner ; that is, on the upper face next the back : at or near the lower right-hand corner, the name of the contained species may be written either with a pencil or in ink. 814. The genera should be arranged in the herbarium accord- ing to some systematic work, and numbered accordingly on the covers. 815. The herbarium must be preserved in close cabinets or cases free from the access of dust. Tin cases, just deep and wide enough to receive comfortably the genus-covers, and about six inches high, the hinged lid being one end, may be recom- mended for a small collection, as they are dust and insect proof, are portable, and may readily be arranged on shelves. But, for any herbarium of considerable size and continued growth, wooden cabinets with well-fitted doors are to be preferred ; the interior of the cabinets being divided into pigeon-hole compart- ments, fully 12 inches wide in the clear and 17 inches deep, and not over 6 inches or in small herbaria not over 4 inches high. Into such pigeon-holes, the genus-covers with their contents will slide readily, and may be compactly stowed awa}-. An index to the genera of each order may be affixed to the interior of the cabinet doors, or pasted upon the upper face of thin boards, inserted at the beginning of each order. The name of the order, written or printed in bold letters, may be pasted upon the front edge of this board, or upon a flap of card-board affixed to it. Moreover, it is well to write the name of the order upon each genus-cover. 816. Except in public collections, where fixed cases may be preferred, the cabinets should individually be small, only three or four feet high, and containing only two or four vertical rows 384 PHYTOGRAPHY. of compartments. Such cabinets can be increased in number as required, arc portable, and can be disposed in any order, side by side or one surmounting another, as ma}- be most convenient. The doors should be so constructed as to oj.cn and shut readily, but to close tightly, so as to exclude du>t and insects.1 1 An excellent plan for small and inexpensive herbarium cabinets, of a portable characiiT, N proposed and illustrated by Dr. Parry, in the American Naturalist, viii. 471. Kadi >inall case is in fact a plain wooden box, wide enough to hold two tiers of pigeon-hole compartments, and of anv desirable height (three compartments high in Dr. I'arry s plan, but double the number might lie better) : the entire front consists of a pair of doors meeting in the centre, there fastened by a rlush spring catch; the doors bevelled on the in>i !e. with a corresponding bevel on the case, to which they are attached bv out- side binges, >o that in opening at a right angle there are no sharp corners to hinder the drawing out of the herbarium papers; also allowing the cases to >tand close side1 by side, as well as one upon another, without interfering with the free opening of the doors. These, moreover, may swing quite back against the' sides without in any way straining the hinges. For lifting, a pair of flush handles, countersunk to the level of the wood, may be attached to the sides. When the herbarium has to be removed to a distant place, these cases, having no projecting knobs or handles, will go readily into ordi- nary packing-boxes. ABBREVIATIONS. L. OF NAMES OF BOTANISTS AND BOTANICAL AUTHORS. Adi. = Acharius. Berkh. = Adans. Adanson. Berken. Afz. Afzelius. Btrland. Ag. Agardh. Bernh. C. Ay. C. A. Agaijdh. Btrt. J. Ag. 1 J. G. Agardh, son. Bertol. Bess. Ait. Alton. Bteb. All. Allioni. Bigel. Amm. Amman. Bisch. Anders. Andersson of Stockholm. Bcehm. Andr. Andrews. Boerh. Andrz. Andrzejowski. Boiss. Aresch. Areschoug. Boland. Arn. Arnott. Bong. Arrh. Arrhenius. Bonpl. Asch. Ascherson. Bark. Aubl. Aublet. Borsz. Brack. Bab. Babington. Brebis. Bail. Baillon. Bref. Bulb. Balbis. Brew. $• \ Baldw. Baldwin. Wats. \ Balf. Balf our. End. Barn. Barne'oud. Brong. Burr. Barrelier. Brot. Bart. Benj. Smith Barton. Bronss. W Bait. Wm- P. C. Barton, nephew. Br.. R. Br. Bartr. John Bartram. P. Br. Bartr. f. Wm. Bartram. Brunf. Bauh. ' Bauhin. Buckl. Beauv. Palisot de Beauvois. Bull. Benj. Benjamin. Burin. Benn. J J. Bennett. Buxb. A. Benn. A. W. Bennett. Bentk. Bentham. Cam. Berg. Bergius. Camb. j Berk. M. J. Berkeley, Cambes. j Berkhey. Berkenhout. Berlandier. Bernhardi. Bertero. Bertoloni. Besser. Marschall vonBieberstein. Jacob Bigelow. Bischoff. Bcehmer. Boerliaave. Boissier. Bolander. Bongard. Bonpland. Borkhausen. Borszcow. Wm. D. Brackenridge. Brebisson. Brefeld. W. H. Brewer & Sereno Watson. Bridel. Brongniart. Brotero. Broussonet. Robert Brown. Patrick Browne. Brunf els. Buckley. Bulliard. Burm an. Buxbaum. Camerarius. !• Cambessedes. 26 ABBKEViATiUNS. ' '" 111 />d. = Campdera. Eat. = Amos Eaton. Cand. DeCandolle, usually DC. D. C. Eat. D. C. Eaton, grandson. < '. Cassini. Edw. Edwards. < 'till A'6. Catesby. Eliren. Ehrunberg. < 'ao. ( ';iv:iliillcs. Ehrh. Ehrhart. < ; ro. Cervantes. Eidd. Eichler. C/tuin. ( 'liamisso. Eiseng. Eisengrein. < '/iiijim. A. \V. Cliapman. Eli. Elliott. ( '/nir. Chavannea. Endl. Endlicher. CliOlS. Choisy. Enyelm. EngL-lmann. Clayt. Clayton. Engl. Engler. ( 'Iu8. ( ln>ius. Esc/is. Ksdischoltz. Col/ad. Colladon. Eschw. Esc) i\v oiler. Colin. Colmeiro. Etttngsh. Etiingshausen. Coin in. ( 'iimmelin. Corn. Cornuti. !•'< ndl. Fendler. Coss. Cosson. Feuil. FeuUl^e, Cunn. Cunningham, A. or J. i'unji rlt. Fingerhutb. < 'lift. Win. Curtis. Fisch. Fischer. M. A. Curt. M. A. Curtis. I-'urs/c. Forsk&l. Forst. Forster. Dalech, Dale-champs. Foam. Fournier. Dalib. Dalibard. Fresen. Fresenius. /><„•/. Darlington. Freyc. Freycmet. DC. ) Fred. Froelicb. DeC. \ A. P. DeCandolle. A. DC. AlphonseDeCandolle.son. Gcpiin. J. Gaertiu-r. Cos. DC. Casimir DeCandolle, tlie Gairtn.f. C. T. (Ja'rtner. Decne. Dccaisne. [grandson. Gardn. Gardner. Deless. 1 )<>lrssert. Gar id. ( iaridel. Demist. Dennstedt. Gasp. ( ias]>arrini. Desc. Descourtilz. Gaud. Gaudin. Desf. Desfontaines. Gaudich. Gaudichand. Desj. Desjardins. Germ . Germain. 1 ' >i smar. Desmazieres. Gesn. Gesner. 1 >< fiinoul. Desmoulins. Gilib. Gilibert. Df-sv. Dcsvaux. Ging. (iingins de Lassaraz. Dicks. Dickson. din. (iisckc1. Diesb. Diesbach. Gled. Gleditsch. 1 >u ter. I >lcii'rich. Gleich. GU-ichen. Dietr. Dietrich. Glox. Gloxin. Dill. Dillenius. Gmel. J. G. Gmelin. Dillw. Dillwyn. C. Gmel. C. C. (iiiH-lin of Baden. Dod. DddoiuiMis (Dodoens). S. Gmel. S. G. Gmelin. D'Orb. D'Orbigny. God,: Godron. Dor st. Doreteniufl. Ga'/i/i. Goeppert. />nin/l. Douglas. Goi'd. ( ioudi'iiougb. Drej. Drejer. Grcn. Grenier. 1 >n/'i/i(l. Dryander. Gr, r. GrevilK'. Dufr. Dufresne. Griseb. Grisebach. Dnliam. Diiliaiiii-l du Monceau. Gram. Greenland. I iiunort. Duniortier. Gron. ) Gronovius. Dun. Dunal. Gronov. ) ABBREVIATIONS. Guett. = Guettard. Guilt. Guibord. Jacq. = Jacq. f. Gui/lem. Guillemin. J. St. Hi (ii/uitp. Guimpel. Jord. Gann. Gunnerus. Jumjh. Guss. Gussone. Juss. Adr. Juss Haqenb. Hagenbach. Hall. Haller. Hani. Hamilton. Kcemp. r\ ti rvt Hunb. Hanbury. Hanst. Hanstein. I \ ' 1 1 Ol> > Kaulf. J\.ll[(Jtl)* Hurtm. Hartmann. Hurtw. Hartweg. Kirschl. Kit Haru. Harvey. ±\. tt, Krelr Hass. Hassall. i \ ' 1 1 f . A'/ij'/A Hassk. Hasskarl. /VO/ ill. Hausm. Hausmann. Kostel. Haw. Haworth. Kremp. Hebens. Hebenstreit. Kromb. Hedw. Hedwig. Kuetz. Hegelm. Hegelinaier. II«/etsc/t. Hegetschweiler. L. Heist. Heister. Labill. Hddr. Heldreich. Laist. Helw. Helwing. Lag. Hemsl. Hemsley. Lull. Henck. Henckel. Lam. Henfr. Henfrey. Lamb. Hensl. Henslow. Lamour. Herb. Herbert. Langsd. Ilerm. Hermann. La Pei/r. Hild. Hildebrand. La Pyl. Hoc/tst. Hochstetter. Ledeb. I I off m. G. F. Hoffmann. Lehm. H. Hoffm. Hermann Hoffmann Lem. flo^vnanns.Hoffmannsegg. Lesq. Hofm. Hofmeister Less. Ho/ie n. Hohen acker. Lestib. Holmsk. Holmskiold. Lev. Iloinb. Hombron. L'Her. Hook. Win. J. Hooker. L'Hcrm. Hook.f. J. D. Hooker, son. Liebin. Hopk. Hopkirk. Lightf. Hornem. Hornemann. Lilij. Hornsch. Hornschuch. Lindh. Horsf. Horsfield. Lindbl. Houst. Houston. Lindcnb. Hontt. Houttuyn. Lindh. H/ids. Hudson. Lindl. Hueb. Huebener. Linn. Humb. Humboldt. Linn. f. HBK \ Humboldt, Bonpland, and Lodd. 1 Kunth. La-fl. 388 ABBREVIATIONS. Lees. = Lceselius. Naud. = Lois. Loiseleur-Delongschamps. Neck. l.mtd. Loudon. Nees or ) . Lour. Loureiro. N.abE. } Lmlw. Ludwig. T. \ees 1 Lu inn. Luinnitzer. Nestl. Lyngb. Lyngbye. Newb. .\i wm. Mac/. Macfadyen. N(£(J(J. Min-i/il. MacGillivray. Nois. Magn. Magnol. [stein. Nord. M. Bieb. Murschall von Bieber- Not. Marsh, Iluinphrcy Marshall. Nutt. Mars. Marsili. Nyl. Mart. Martius. Nym. Mass. Massalongo. Musi. Masters. (Ed. Maxim. Maxiinowicz. (Erst. Med. Medikus or Medicus. Oliv. Meisn. 1 Meisner or Meissner. D. Oliv. M: issn. } Orb. Mt ucg. Meneghini. Orph. M, nz. Menzies. On. Mir/. Mertens. Oudem. Mi //en. Mettenius. Mich. Mielieli. P.deBeauc Michx. MX. | Andre' Michaux. Pall. Panz. Michx. f. F. A. Michaux, son. Park. Midden. Middendorff. Purl. MM. Philip Miller. Pasq. Mill. J. John S. Mueller or Miller. Pao. Mi,/. Miquel. Perl. Mirb. Mirliel. Pers. Mitch. John Mitchell. Philib. Mitt. Mitten. Planch. Mm:. Mocino. G. Planch. Mnlk. Molkenboer. Pluk. Mont. C. Montagnc. Plum. Mm/. Moquin-Tandon. Pp. .1 loric. Moricand. Pair. Moris. Morison. Poit. Murr. Morren. Poll. M< "nj Mougeot. Post. Mm //..1/v/. .1. Mueller of Arpau. Pourr. /•'. Mn, II Ferdinand Mueller. Pringsh. 0. .I/,/,// otto Mueller of Denmark. Pritz. Muhl. Muhlenberg. Putt, r. Muni M nnling. Murr. J A. Murray. Hnhrnfi. A. Murr Andrew Murray. JiaJIk. Raf. Nacc. Naccari. Jinsp. Nag. Na-geli. Red. Naudin. Necker. C. F. Nees von Esenbeck. T.F.L. Nees von Esenbeck. Ni .-tier. Newberry. Newman. Nceggerath. Noisette. Nordstedt. Notaris. Nuttall. Nylander. Nyman. CEder. CErsted. Olivier. D. Oliver. A. or C. d'Orbigny. Orphanides. Ortega. Oudemans. .Palisot de Beauvois. Pallas. Panzer. Parkinson. Parlatore. Pasquale. Pa von. Perleb. Pcrsoon. Philibert. J. E. Planchon. Gustave Planchon. Plukcnet. Plumier, Lat. Plumerius. Poeppig. Poiret. Poiteau. Pollich. Postcls. Pour ret. Pringshcim. Pritzcl. Putterlich. Ixahenhorst. Radlkofer. Rafinesque-Schmaltz. Raspail. Redoutc. ABBREVIATIONS. 389 Reich. = Reichard. Scop. = Scopoli. Reichenb. H. G. L. Reichenbach. Seem. Seemann. Reichenb.f. H. G. Reichenbach, son. Sendt. Sendtner. Reirnv. Reinvvardt. Seneb. Senebier. Reiss. Reisseek. Ser. Seringe. Retz. Retzius. Stub. Seubert. Reut. Reuter. Siith. SibtUurp. Rich. L. C. Richard. Sieb. Sieber. l^V[ \ Achille Richard. A. Rich. ) Sieb. Soland. Siebold. Solander. Ric/iards. John Richardson. Sow. Sowerby. Richt. Richter. Spenn. Spenner. Ridd. Riddell. Spreng. Sprengel. Rio. Rivinus. Sternb. Sternberg. Rcelil. Roehling. Steud. Steudel. Rcem. J. J. Rcemer. Stev. Steven. M. J. RamM. J. Roeiner. Sull. Sullivant. BTn' & \ Rcemer & Schultes. Sen. ) Sw. Swartz. Rcep. Rceper. Tare,. Targioni-Tozetti. Rohrb. Rohrbach. Ten. Tenore. Rostk. Rostkovius. Thorns. Thomas Thomson. Rothr. Rothrock. Thuill. Thuillier. Rottb. Rottboell. Thunb. Thunberg. Rottl. Rottler. Thurb. Thurber. Roum. Rouniegere. Thurm. Thurraan. Roxb. Roxburgh. Tod. Todaro. Roy. Royen. Torr. Torrey. Rudb. Rudbeck. Torr. Sf Gr .Torrey & A. Gray. Rupr. Ruprecht. Tourn. Tournefort. Tratt. Trattinick. Sacc. Saccardo. Traut. Trautvetter. Sadl. Sadler. Trev. , Treviranus. St. Hil. A. Saint-Hilaire. Trin. Trinius. Salisb. Salisbury. Tuck. Tuckerman. Salm-Dyck.Prmce Jos. Salm-Riffer- Titrcz. Turczaninow. Sauss. Saussure. [schied-Dyck. Turn. Turner. Schimp. Schimper. Turp. Turpin. Sr/iL Schkuhr. Sclilecht. Schlechtendal. Vaill. Vaillant. Sc/i/eich. Schleicher. VeiU. Veillard or Vieillard. Schomb. Schomburgh. Vauch. Vaucher. Schrad. Schrader. Vent. Ventenat. Schreb. Schreber. Vill. Villars, or Villar. Schueb. Schuebeler. Vis. Visiani. Srhult. Schultes. Vittad. Vittadini. Schultz ) 0. H. Schultz, Bipontinus Viv. Viviani. Bip. } (Zweibrucken). Vog. T. Vogel. Schiim. Schumacher. Schnitd. Schnitzlein. Wahl. Wahlenberg. Schiccegr. Schwaegrichen. Wahlst. Wahlstedt. Schwein. Schweinitz. Watts. Wnldstein- Schweiiif. Schweinfurth. Wall. Wallich. Schwend. Schwendener. Wallm. Wallman. ABBREVIATIONS. \Vullr. = Walp. Wait. II Wats. H.C.Wats S. Hate. Web. Wedd. II i in m. Welw. II 'ruder. Wendl. WHes. Wallroth. Walpers. Walter. Wangenheim. Wanning. 1'. W. Watson. H. C. Watson. Sereno Watson. Weber. Weddell. We imiiann. Welwitscli. \Vciuleroth. Wi-ndland. Wikstrom. \Vil"-im. or l>i,-m. Decimetre. /'»Mr. Description. Diff. Differentiae, the distinguishing marks. Ed. Edition. r.mlir. Embryo. /.'-N. Essential, as Char. Ess. / ••!. Excluding, or being excluded. Eicl. Sy». Excluding the synonym or synonyms. Fam. Family. Fil. Filament of the stamen. Fl. Flower (flos) ; Flora, or some- times Floret, it flowers. Fcem. Female plant, flower, &c. Fol. Folium, leaf. Fr. Fruit. Frm-tif. Fructification. Gen. Genus or Generic. Germ. Germen, Linnaean name for ovary ; also Germination. //. Herbarium. Hal. Habitat, place of growth; sometimes for Habeo, I have. Hi rh. Herbarium. 7/iirf. Ilortus, garden. Hortul. Hortulanorum, of the gar- deners. /r. Icon, a plate or figure. ///. Illustris, illustrious. fun/. Unpublished. I >il'. Inferior. Infl. Inflorescence. ////•. Involucre. Lot. Lateral, or relating to width. Lin. Linea, a line (the 12th of an inch). Lit., Litt. In a letter or letters. /. c. Loco citato, in the place cited. ABBREVIATIONS. 391 Masc. Male plant, flower, &c. Mill, or mm. Millimetre. Mss. Manuscripts. Mus. Museum. jV. or No. Number. Nat. Natural. Xttm. Nomen, name. Obs. Observation. Ord. Order. Oo. Ovary. />. Page, or sometimes Part. Ped. Peduncle or Pedicel, or Pedalis, a foot long or high. Peric. Pericarp. Perig. Perigonium. Pet. Petal or Petiole. Plst. Pistil. Plac. Placenta. Poll. Pollicaris, an inch long. p. p. Pro parte, in part. Prodr. or Prod. Prodromus. Rad. Radix, root ; or Radical. Ram. Ramus, branch. s. Seu, or Sive, Latin for or. Sect. Section. Segm. Segment. Sem. Semen, seed. for Sep. Sepal. Her. Series. Sice. Siccatus or Siccus, dried or dry. Spec, or Sp. Species, or specimen. Spont. Spontaneous. Stum. Stamen or is laminate. Sup. Superior. Si/n. Synonym or Synopsis. T. or Tab. Tabula, plate. T. Tom us, volume. V. Volume: sometimes for Vel, or; sometimes Vide, see. Var. Variety. Veg. Vegetation, characters of Vern. Vernal. v. s. Visa sicca, or Vidi siccam. v. v. Visa viva, or Vidi vivam ; the first indicating that a dried speci- men of the plant, the second that the living plant has been exam- ined. v. s. c. and v. s. s., indicates that the dried specimen was cultivated (c) or spontaneous (s). v. v. c. and v. v. s., that the living plant seen was cultivated (c) or spontaneous (s). SIGNS. 1. SIGNS USED BY LINNAEUS. 0 An annual plant. $ A biennial. 11 A perennial, b A tree or shrub. * Affixed to a reference, means that a good description will be found there, f Indicates an obscure or doubtful species. 2. SIGNS USED BY DECANDOLLE AND LATER WRITERS. 0 A monocarpic plant, i. e. which dies after once flowering and fruiting, either annual or biennial, or of longer duration. W Annual. (?) Biennial. (°°) Monocarpic perennial, such as Agave. 392 SIGNS. 11 Perennial herb. •^ Suffrutex, an undcrshrub. 5 Frutejc, a shrub. 5 Arbuscula, a tree-like shrub of ten to twenty-five feet in height. 5 Arbor, a tree. <~* A climbing plant. A An evergreen. $ Male plant or flower. ^> Female plant or flower. T£ Hermaphrodite plant or flower. oo Indefinitely numerous, e. y. ao-andra, polyandrous. ? A sign of doubt. " Thulictrum ? Jajionicum," doubts if the plant is really a Thalictrum. " Thulictrum Juponicnm, Thunb.?" doubts if the plant in hand is truly the species of Thunberg. Thalictrum Japoni- cum, Thunb., Willd.? doubts whether Willdenow's T. Japonicum is really tliat of Thunberg. ! A sign of certainty. As " Thulictrum anemonoides, Michx. ! Fl. Bor. Am. p. 322," as used by DeCandolle, affirms that he lias seen an authentic original specimen of this author. Affixed to the name of a collector, as " Virginia, Clayton ! " it affirms that the writer has examined a specimen collected by the person to whose name it is appended. Between two figures, as in " Stamens 5-10," indicates the extremes of difference, as that the stamens are from five to ten. 0 ' " The signs for degrees, minutes, and seconds, as 1°, 2', 3", are used in Gray's Manual of Botany of the Northern United States, for feet (°), inches ('), and lines (")• With European authors, usually the sign for minutes is for feet ; that of seconds for inches : thus 1', a foot high ; 1", an inch long; and 1"', a line long. O— Cotyledons accumbent to the radicle. Oil Cotyledons incumbent on the radicle. GLOSSARY, OR DICTIONARY OF BOTANICAL TERMS, ENGLISH AND LATIN, COMBINED WITH AN INDEX. THIS Glossary is intended to contain all the principal technical terms (substan- tive as well as adjective) of structural and systematic Botany, as far at least as concerns Ph.tnogamous plants. Most of the special terms relating to the lower Cryptogamia and to Vegetable Anatomy and Physiology are relegated to the vol- umes devoted to those departments. The annexed numbers refer to pages of this volume. Very many of the terms are seldom employed, or are wholly out of use. The principal Latin terms are given separately only when there is no English equiv- alent differing merely in the termination. When the word is essentially the same, the Latin termination (of adjectives in the nominative masculine only) is annexed to the English word in a parenthesis. The changed termination goes back mostly to the penultimate consonant. It is unnecessary in a work like this to accentuate all the technical words; but, in the case of words liable to mispronunciation, an accent-mark is placed over the syllable which takes the principal accent. The glossary, as here drawn up, may serve to indicate the meaning of the commoner descriptive specific names of plants. A, privative, as the initial in many words of Greek derivation, signifies the absence of the organ mentioned*, as, ylpetalous, destitute of petals ; Aphyllous, leafless. In words be- ginning with a vowel, this prefix is changed to an; as, --Ireanthous, flow- erless ; ^wantherous, antherless. Abbreviations, 385. Aberrant. Wandering, applied to spe- cies, genera, &c., which differ in some respect from the usual or normal char- acter of the group they belong to. Abnormal (Abnormis). Differing from the normal or usual structure. Aboriginal. Strictly native; indigenous. Abortion (Abortus). Imperfect develop- ment or non-development of an organ ; 179, 187. Abortive (-ivus). Defective or barren. Abrupt (Abruptus). Terminating sud- denly ; the opposite of tapering. Abruptly pinnate. Pinnate without a terminal leaflet or appendage ; 101. Acant/tocladous (-us). Having spinjl branches. Acanthophorous (-us). Spine-bearing. Acaulescent (-ens). Stemless, or appar- ently so, with no proper caulis; 45. Acaulls. Stemless; same as Acaulescent. Accessory. Something additional, or of the nature of appendage. Accessory Bud*. 44. Accessory Fruits, 300. Accrescent (-ens). Increasing in size with age, as often occurs with the calyx after flowering. Accrete (-us). Grown together, or con- solidated with some contiguous body. Accumbent (-ens). Lying against an- other body. Accumbent Cotyledons. With edges against the radicle: 313. Acephalous (-us). Headless. 394 GLOSSARY. Jlcerose (-<&«). Needle-shaped, like the leave-, (it 1'ines. Acetdbuliform (-W/H <'.<). In the form of a shallow open cup or saucer. Achcenium or Achenium. A small, drv and hard, one-celled, one-seeded, inde- hiscent fruit; strictly one of a single and iree car])d; but extended to simi- lar lines of more than one carpel, and also with adnate calyx; '2!»4. (Achce- nium is etymologically ihe proper orthography; but m-l,, niinn is be- coming tlie commoner form.) Ackcenocarp (-arpium). Cencral name of a dry and indehiscent iruit ; L'.'J. Achenodium, Such a double aclienium as that of I'mbellifenc ; a Cremo- carp. Achlamydi mis (-eus). Destitute of peri- anth ; l!ll. A bristle. /ni- (-nria). r.ristle-shaped, or slen- der needle-shaped. Acindciform (-«/•/»/.<). Seyniitar-shaped; curved with rounded point, tliicker on tlie strain-liter edge than on the con- vex edge. jii-iin>. a regular symmetrical (lower; 175. Aciitiii.-i-iiliif. Somewhat acute; acutish. A/l< /jtliima (-ii.-i. Adelphi, brothers). Sta- mens with ccalesceiit or clustered li la- ments are monadelphous, diailelplious, &c., according to the number ot ' A*il/>/ti .<. Air-plants; 35. Verdicpris-colored. i -ii/i.*). l.'elating to summer. tiiin (-in}. The disposition of the parts of a flower in the bud ; 132. GLOSSARY. 395 o. A form of aggregate fruit ; 300. sfctheogamia, ^Ethoyamous, 340. Aj/iiiity. True and near relationship; •\-27, 330. Agamous or Agamic. Destitute of sexes. I Heaped or crowded Agglomerate. ) . < into a dense cluster, A(/gregate(-atus). I . \ but not cohering. Aggregate Fruits. Those formed of aggregate carpels of the same flower; 298, 301. Ayrtstis. Growing in fields. Air-plants. Plants unconnected with the ground; 35. Akene, Akenium. See Achsenium. Ala (pi. tarch. Analoi/y (Analotjia). Likeness in cer- tain respects. As distinguished from a (fin it i/, it means resemblance in cer- tain respects only, not in the plan of structure. Thus, a Ranunculus is analogous to a Potentilla. but there is no near affinity or relatio: ship be- tween the two. And the tendril of a Pea, that of a Smilax, and that of the Grape-vine are mvilogues; i. e , are analogous organs, but are not linitto- loi/ues; for the first answers to a leaf. the secouJ to stipules, and the third 396 GLOSSARY. to a stem. The spur of a Larkspur is analogous to one of the five spurs of Columbine, but not homologous with it ; for the first is a sepal, and the second a petal. Aiiniitli-iiit.i. Institute of stamens. Aii(inther(jus(-us). Destitute of anthers. .-i <-//.<}. !• lowerless. The connection of veins, £c., by cross-veins, forming reticu- lation. iiii/i-njiiiiiii (-us), wrongly Anatropal. Tlie reversed ovule, with mieropyle close hv the side of the hilmn, and V chala/.a ;it the opposite end; '27\>. F.ngl. Aui-'ijittal. Botanically always used in the sense of two- edged. Am/1 /•, iiiii/ni, iinilrnm. In Greek com- pounds, the male. Anilro-ilurciuus. With flowers on one plant hermaphrodite, and on another stamiiiute only; 191. Andriecium. The stamens of a flower coll.-etivcly; Hi,r), 24!». Ainli-uijynous (-us). Said of an inflores- cence composed of both male and female flowers. t&ndrophore (Andrdphorum). A sup- port or column on which stamens are raised. Androus. See Ander. Anemdphilota. Literally wind-loving. Said of flowers which are fecundated by wind-borne pollen; 217. Anfractuosus. Abruptly bent hither and thither, as the stamens of Cucur- bita. Angiocarpotu (-us). When a fruit is covered by some envelope. Aiif/ivf/ii riniii. A Limuean artificial order; 337. Anffiospermous, Angiotperma, Anyio- .ipi-rms. Plants with seeds borne in a pcr'u-arp; 25:1. Aiii/iilnr Dircryence of leaves; 123. Aiiifiniirnius (-us). Une(|iia! in number in the different circles of the flower; u asymmetrical. Ani.to/1' /«/«//< (-HH). With unequal petals. An'uophylloua (-us). Unequal-leaved: i. e., the two leaves of a |>air unequal. A ii 1st '.-••/ 1 minimi* (-iif). When the >ta- mens are not of the number of the petals. Antwtinus. A year old, or in yearly growths. Annual (Annuns). Of only one year's duration ; :!(). Annular (-am). In the form of a ring; or marked transversely by rings. The latter more properly. Anitulnti (-ntus). Marked with rings. Jliinulus. A ring, such as that with which the sporangia of Mime Ferns and Mosses are furni>hcd. Anupliytes (Anuphytu). Name of group comprising Mosses, &r. Auli i«:.r, as to position, denotes the front side, or averx- from the axis of intio- iv-cence; Itiit. Anthela. A deliijuocent and paniculate cyme, with median ramilication, and the lateral axes overtopping the central, as in Juncus tennis, &c. May be either a Drepanium or a Khi- pidium. Anthemy, Antkemia. A flower-cluster of any kind ; 144. Anther (Antln'rn). The polliniferous part of a stamen; 105, 251. Antltt ridiinii. All analogue of the an- ther in Cryptogams. Anthi rij'< i-ii/i.t (-//.<). Anther-bearing. Jlntlusi.t. r|'h,. time ;it which a flower is perfected and opens; or the act of expansion ol' a tinucr. Anthoc(i>'pou$( -«.iirli«tc (-atus). Bearded; beset with lung and weak hairs. Barbi-llntv (-it/ us). Beset with- shorter and stiti'.'r hairs or linrhvlUe. Hartivllultitt: (-ulus). Uiiuintitive of the preceding. Buik. Tlie rind or cortical portion of a stem, especially of an exogen; 76. Bnxul ( lin.-i'ilni-'it!. Kelating to tlie base. Busid-n, roed. \\"ith nerves all from the base of the leaf; 92. ll'ift ( li'itif). The extremity by which an organ is attached to its support. lltitiiltn . Cells of the fructification of .Mushrooms which bear the spores. Btutijifi-d (-us). Attached by the base or lower end ; 2.>i. />ii.i-), two, twice, or doubly. Biaritiiiiinite (-ense for inflated. Bursicula. A small pouch (bursa); such as that which encloses the disk or gland of I lie eaudicle of the pollin- iniii of an ( )rdiis. /ini-si, -it/, i/ns. Fnrni.-hed with a bursi- cula or pouch. /•' iceotu (-(».<). Composed of fine threads, like bi/f.iiin or line flax. iis (-»s). Dropping off very early, as the calyx of a Poppy at the time of expansion : '243. Cceruleus. Sky blue, or pure blue. Ccesius. Lavender-color ; pale green with whitish or gray. Ciiliit/tl'Jiiim, Ciil'ithis. Literally a bas- ket ; a name for the head of flowers (or better for the involucre only) of L'om- positse Calathiform (-vrmis). Cup-shaped ; of somewhat hcmi-plierical outline. Calcar. A spur; mostly used for the nec- tariferous one of a calyx or corolla. Cdlcarate (-tifus). Furnished or pro- duced into a spur. Cdlceulate (-iitus), or Ctilceiformis. Shaped like a slipper or shoe. Callose (-osus). Bearing callosities (calli), or hard protuberances. Calctis. Bald, as an akene without pappus. Calycdnthemy. Name of the monstros- ity in which the calyx imitates an exterior corolla ; J74. Calyciflorotu (Calyciflorce), 340. Calycine ( < 'n/i/rnuis). Kelating to calyx. Culyculate (-atus). Bearing bracts next the calyx which imitate an external or accessory calvx. Calyculus. An involucre or involucel imitating an additional calyx. Culyptra. The hood or veil of the spore-case of a Moss; or some cover- ing body like it. Calyptrnte (-ntns). Furnished with a calyptra, or .something like it. Calyptriform (-oi-mi's). Calyptra-shaped; as the cal}-x of F.schscholtxia. Calyx. The flower-cup, the exterior perianth ; 164. CdiiKira and its diminutive Crnn-'rula (chamber) are sometimes used for the cells of a fruit. Cambium. Old name of the viscid mat- ter between bark and wood in com- mon trees or shrubs in spring; now used for the nascent structure there forming, or Cambium Injcr; 78. Cninji.ii/iilntr (-ntim). Bell-shaped ; elon- gated cup-shaped or shorter, and broad from the base ; 249. Campaniformis. Same a- (' unpannlate. Campylospermmw (-».<). Curved-seeded. Said of seed-like fruits or carpels, as those of some I'mbellifenv, in which the contained >eed i- involute by the lateral edges. -o a- to produce a longi- tudinal furrow on the ventral face. Campyldtropota (-«.<), or less correctly Campylotropal, or Campulitropous. An ovule or seed which is curved in its formation so as to bring the GLOSSARY. 401 micropyle or true apex down near to the hilum; 279. Canalicaulate (-atus). Channelled, or with a longitudinal groove. Cdncellate (-dtus). Latticed; resembling lattice-work. Candid us. Pure white. Canescens. Hoary, usually with gray pubescence. Canus. Gray-white; whiter than the preceding. \ So slender that it Capillaceous(-eus). f may be compared Capillary (-aris). I with the hairs of ) animals. Capitate (-a(us). Head-shaped, or col- lected in a head; 147. Capitellate (-atus). Diminutive of Cap- itate. Capitulum. A head of or simple globu- lar cluster of sessile flowers; 147. Caprcolate (-atus). Bearing a tendril (capreolus). Capsule (-ula). A dry and dehiscent pericarp composed of more than one carpel ; 289, 293. Capsular. Of the nature of, or relating to, a capsule. Capsultferous. Capsule-bearing. Carci'i-ulus. An unused name for an indehiscent and several-celled dry fruit; 297. Carina. A keel; used either for the two combined lower petals of a papil- ionaceous corolla (185); or for a sa- lient longitudinal projection on the centre of the lower face of an organ, as on the glumes of many Grasses. Connate {-atus). Keeled. Cat-iopsis or Carynpsis. A grain ; a seed- like fruit with thin pericarp adnate to the contained seed ; 295. Carneus. Flesh-colored, very pale red. Caro. Flesh, as the pulp of a melon, or the fleshy part of a drupe. Carpndilium. Synonym of Cremocarp. Carpel, Carp^lluin. A simple pistil, or an element of a compound pistil, an- swering to one leaf; 167, 260. Carpid, Carpidium. Synonym of carpel. Cnrpolngy. The botany of fruits. Carpophore (Carj>6/>/i<>rum). A portion of receptacle prolonged between the carpels; 212. Cdrpophyll (Carpophyllum). Literally fruit-leaf; synonym of Carpel ; 260. Cartilaginous or Cnrtilagineous (-eus). Of the texture of cartilage or gristle; firm and tough. Caruncle (Caruncula). An excrescence at or about the hilum of certain seeds; 308. Caryophyllaceous(-eus). Resembling or relating to the corolla of Dianthus Caryophyllus (246), or to the Pink family. Caryopnis. See Cariopsis. Cngside «•»•. Helmet-shaped. Cassus. Empty, as an anther contain- ing no pollen. Cntitrate (-atus). Said of a stamen which wants the anther. Catapetalous (-us). Where petals are united only by cohesion with united stamens, as in Mallow. Cataphylla. Answers to the German " Niederblatter," or under-leaves, those at the beginning of a growth, cotyledons, bud-scales, scales on rhi- zomes, &c. ; 6. Catenulate (-dtus). Formed of parts united end to end, like the links of a chain. Catkin. A scaly spike (see Ament); 150. Caudate (-atus). Furnished with a tail (cauda), or with a slender tip or ap- pendage resembling a tail. Caudex. A trunk or stock of a plant ; 50. Caudicle (Caudicula). The stalk of a pollinium, £c. Caulescent (-ens). Having an obvious stem. Caulicle (Cauliculus). The initial stem in an embryo, generally named the Radicle; 10." Cauline (-inns). Belonging to the stem. Caulis. Greek form Caulun. The stem of a plant. Caulocdrpic or Caulocarpous. Applied to plants which live to flower and fructify more than once or indefi- nitely. Caulome, Cauloma. The stem-part of a plant. Cephalanthium. One of the names of the head or capitulum in Composite ; 148. Cell ( Ctllula). The anatomical element of plants; 28. The cavity of an anther which contains the pollen, or an anther-lobe, thus taken in the sense of the circumscribing wall as well as the cavity; 251, 254. The cavity, or any one cavity of an ovary or pericarp, containing the ovules or «feds; 262. Cellular Plants, Cellulares, 340. Cellule (-ula). Diminutive of cell; of 402 GLOSSARY. the same meaning as Cell in vegetable anatomy : -JS. Cellulose, 'lli.' material, chemically con- sidered, of which the wall of the cell consists. Cenobium. \ name of the peculiar four- parted fruit (or the four nutlets around a i ,,111111011 style') whii-h distinguishes l.;i!ii;it;i' and l!orraginace;i-. <'< nt 1 1 '/'//(/"/. Tending or developing from the centre outward. I ', iiii-iii< inl. 'rending or developing from without toward the centre. Cephalanthium. Synonym of Antho- diiiiii. Ceriiiiiiin. \ silii|uiform capsule, such as that of <'nrydalis, (.'Iconic, &c. Cereal. Belonging to corn and the allied grains. Cn-inus. Of the color of wax. Cernuous (-uus), Nodding. Chu'in. Greek for a brittle, Latin Seta. Clinji'. Small scales; dry and depau- perate liracts; such as those on the re- ceptaclc of a siintlower and many other Composite ; al>o glumes of Grasses. Chaff ij. Provided with or having the icxinre of chaff. Chnluzn. The part of an ovule where coats and nucleus are confluent; '277. Channelled. Hollowed out longitudi- nally like a gutter. See Canaliculate. f/irirm-tir. A diagnostic description, or the enumeration of essential differ- ences ; 3iii. Chasmdyamy. The opening of the peri- anth at (lowering time; the opposite of Cleistogamy. fliitrt,i<-,«ii* (-ens). Having the texture of writing-paper. Chlorophyll. The green matter of leaves and other vegetation ; 76, 88. Chloros. G reek for green. Kilters into compounds, such as ('li/onnitlius, green-flowered, c/tli>r«>itl/t/, same as I'lilorosis, as when petals turn green; 172. Cklordsis. Literally becoming green, as soini' lloucrs in retrograde metamor- phosis. Also used contrariwise for the loss of a normal green color: 172. fl,,n;lii 1'itt'ilt n-if. \ line of tissue reach- 'ing from stigma to ovary. CkoripetaloUS (-".-•). Same as I'nlvpeta- lous, ;. e. pi'tals unconnected : •_' I I. Chorisepakws. Same as I'oh-xepalous.&c. f/mrixis. Tin- separation of a leaf or phylliim into more than one ; 202. Choristophylltu. Separate-leaved. Chromule (-ula). Coloring matter of plants other than chlorophyll, espe- cially that of petals. fin-i/mif. Greek for golden, or golden- yi-llow; as Cliriifiittiiiis. Yellow-flowered, £c. ' ••//... i ii-Htt-icntii. A scar left bv the fall of a leaf or other organ. Cilin/c (-«/«.-•), i'ili«r!.<. Marginally fringed with hairs. ("t'lniin, pi. ci/in. Marginal hairs, form- ing a trmge, like the eyelash. (The name has In-ni extended 111 scientific books to undividual hairs, and of a surface as well a- edge Cincinnus. A curl : name of a uniparous scorpioid cyme, wliich is Cincinnal ; L56, 157. ( "ni' in-ltijma. Laticiferous tissue. Cinerascens, ('/«< /•«(•< uf. Ash-grayish. Cinereous (-eus). A>h-gray. Cirmabarinus. Cinuabar-i-olor ; scarlet touched with orange Circinul (-(dig). Involute from the tip into a coil; 133. Circinate, or Circinnate (-l,i/lt'i. liranches as- suming the form and function of foli- age : li.'i. 66. flu nit, (-utitx), Cliu-tfurmis. Club- shaped. ff'ir, /fnt,'. Diminutive of Clavate. Clavlculate(-atus). Kurnished \\-\t\\clav- /,-///, i .- viz , tendrils, hooks, or other appliances for climliiug. r/,rw. :;iT). Classification, 315. fliitlirnt, (-utiit). Latticed. f/nif. The narrowed lia-c ,,r stalk which some petaN, \'c., possess: 245. ( 'I, istdgamy, < '/< hti>uirs around the supporting axis; 108. Connective (-iviiiu). A portion of a sta- men which connects the two cells or lobes of an anther; 251. Connivent (-ens). Coming into contact or converging. Conocarpium. An unused name for an aggregate fruit, such as a strawberry, consisting of many carpels on a coni- cal receptacle; 2D8. Consolidated. When unlike parts arc coherent. Continuous. The reverse of articulated or interrupted. Contorted (-us). Twisted; or bent or I\\ i-'ed on itself. In .Estivation, the same as Convolute: 138. Contortuplicnte (-ntus). Twisted and plaited or folded. Contracted. Mil her narrowed or short- ened. Contrary (-ariiis). Opposite in direction to the part compared with : as a silicle compressed eontrarv to the dissepi- ment. Convolute (-11 tin) or Convolutive (-irns). li'olled up from the sides or longitudi- nally. In /Estivation, 138. In Ver- nation, 133. CArnliiiiii ('(-tig). Coral-like. Cdrcuhim. Old name for the embryo, or Cor win in is; 311. Cordate (-atus), sometimes Cordiform (-l tin texture of horn. Coniii'iil i/i (-atus). F'urnished with a little horn. Cornu. A horn ; /. f. a horn-like process; sometimes used for Calcnr, a spur. Cornute(-utus). Furnished with a horn- like process or spur. Corolla. The interior perianth, com- posed of petals; 105, 243. Corollaceous (-eus), Corollinus. Pertain- ing to, or resembling corolla. Corolliferous (-us). Bearing a corolla. i 'on Uijluroiif, ( 'iiruHijIiirce, 340. Coro/lula. Diminutive of corolla. Corona. A crown : an inner appendage to a petal, or to the throat of a corolla ; 210,246. Or any coronet-like append- age at the summit of (crowning) an organ. Cdronntt (-atus). Crowned, having a corona, &c. Coroniform (-ormis). Shaped like a crown or coronet. CSmnjiitt (-atus or -a(ivus). Wrinkled or in folds ; 133. ' 'urtt. r. Kind or bark. Cortirn! (-tills). Kelating to bark. Corticate (-atus). Coated with a bark or with an accessory bark-like cover- ing. Corymb ( Corymbus). A flat-topped or merely convex ami open flower-cluster of the indeterminate or centripetal order; 140. Coi-ymbiferous (-us). Rearing corymbs. Ci'iri/iiibose. In corvmbs. or in the man- ner of a corymb. The corymb of Linmvus and of other writers down to Kirper included most cymes. So that much cymose inflorescence is in descriptions loosely said to be corvmhose, or a stem is said to be corymliosely branched, even when -the evolution is centrifugal ; 140. Costa. A rib ; when single, a midrib or mid-nerve. GLOSSARY. 405 Costal-nerved. With nerves springing from a midrib; 92. Costate (-atus). Ribbed; furnished with one or more longitudinal primary veins or ribs. Cotyledons (Cotyledon, pi. Cotyledones). The " seed-lobes," being the leaves or first leaves of the embryo; viz., the one, or the pair, or rarely the whorl of leaves borne by the radicle or caulicle ; 10, .311, 313. Cotyliformis. Dish-shaped, or wheel- shaped with an erect or ascending border. Crateriform (-ormis). In the shape of a goblet or cup, of hemispherical con- tour or more shallow; 248. Cr&mocarp ( Crenwcdrpium). A dry and seed-like fruit, composed of two one- seeded carpels, invested by an epigy- nous calyx, and separating at matu- rity; 297. Creeping. Running along or under ground and rooting; 53. Crena, Crenatura. A rounded tooth or notch. Cri-nate (-atus). Toothed by crenatures ; scalloped; 98. Crenel, Crenelled. Same as Crenature and Crenate. Cre'nulate (-atus). Diminutive of Cre- nate, i. e. with small crenatures. Crested. Furnished with any elevated line, ridge, or conspicuous elevation on the surface, especially such as may be likened to the crest of a helmet. Cretaceus. Chalk-white; chalky. Cribrose (-osus) and Cribriform (-ormis). Pierced like a sieve. Cribriform Cells, 77. Crinitus. Bearded with long and weak hairs. Critpatus. Curled or crispy. Cristate (-atus). Crested. Croceus, Crocatus. Saffron-colored, i. e. deep reddish-yellow. Cross-breeds. The progenv of interbred varieties; 321. Cross-fertilization. Fecundation by pol- len of another flower and of another individual; 216. Crown. See Corona, 210; 246. Crowned. See Coronate. Crowning ( Coronans). Borne on the summit of an organ. Cruciate (-atus), Cruciform (-ormis). Cross-shaped. Cruciferous (-us). Cross-bearing; used in the sense of Cruciform ; as the "cruciferous" corolla of the order Crucifersc; 246. Crumpled. See Corrugate. Crustaceans (-us). Of hard and brittle texture. Cryptos. Greek for concealed ; whence Cryptogamia. Cryptogamous or Cryp- togamic plants ; 3, 335, 344. Cryptogamous. Pertaining to the above. Cucullnte (-atus), Cucutldris, Cuculli- furrnis. Hooded, or hood-shaped, cowled. Culm ( Culmus). The peculiar stem or straw of Grain-plants and Grasses; 50. Cultrate (-atus), Cultr if ormis. Shaped like a broad knife-blade. Cuneate (Cuneatus), Cuneiform (-ormis). Wedge-shaped ; triangular with an acute angle downward ; 95. Cup-shaped. In the form of a drinking- cup. Cupule (Cupula). The acorn-cup and the like; 230. Cupularis, Cupulatus. Furnished with or subtended by a cupule or any re- sembling body. Cupuliferous (-us). Cupule-bearing. Ciirvinerved (-ius). When the ribs of a leaf are curved in their course ; 92. Cuniserial. In curved or oblique ranks; 124. Cushion. The enlargement at or be- neath the insertion of many leaves. Cuspidate (-atus). Tipped with a Cusp, or sharp and rigid point ; 97. Cut. Same as incised, or in a general sense as cleft. Cuticle ( Cuticula). The outermost skin or pellicle. Cutting. A severed portion of a plant used for bud-propagation ; 43. Cydneus. A clear bright blue. Ci/dfhiform (-ormis). Cup-shaped; in the form of a Cydihus. A drinking-cup, such as a goblet or wine-glass. Cyde. A circle. Sometimes used for one turn of a helix or spire; 122. Cyclical. Relating to a cycle; or coiled into a circle; 119, 120." Cylindraceous. Somewhat or nearly cylindrical. Cylindrical (-us). Elongated and with circular cross-section ; in the form of a cylinder. Cymbceform or Cymbiform (-ormis). Boat-shaped. Cyme (Cyma). A flower-cluster of the determinate or centrifugal type. -KM; GLOSSARY. especially a broad and flattish one; 151. < ijinn-botryose. When cymes are ar- ranged in botryoM' manner; 159. Cynwse (-osus). Bearing cymes, or re- lating to a cyme ; 151. C,/iniil, ( < 'yni nl,i ). Dimiiuit ive cyme, or a portion of a cyme; 151. t'l/iinri'lniiiiiiiii. Name of such a fruit as that of the Hose; tleshy, hollow, and enclosing achenia. C'yjixfla. Name of an achei ium in- vested by an adnatc calyx, as the fruit of Composite ; -'•>•< Ci'/.-itrHtlt. OIK- of the mineral and usually partly crystalline concretions of the ce'ls of the epidermis of or subjacent tissue ol the leal in various plant>, especially in L'nicacca-. Cytolil'ist. An obsolete name for the nucleus of a cell of cellular tissue. Ddct ylose (-osus). Fingered, or finger- shaped. l> i.-i/p/.yllaus (-us). Woolly-leaved. I >, nib, IK (-atus). Whitened over (as if \\ hitewashcd) with a white powder or minute pubescence. Deca. Greek for ten, compounded with various words, such as Dec'iyynia. One of the Linnsean artificial orders; 337. Decayynous (-us). With ten styles or carpels. />, cdm< mus(-us). Often members; 176. />, r • 1/1,1,-i'i. \ Linnaean class with ten stamens; 334. /i,,,i,,,/rous (-us). With ten stamens; 249. Dec.apeta.lous (-us), Decasepalous, &c. With ten petals or sepals, &c. DecliliK'Hf (-«.<). Falling, or subject to fall in season, a* petals after anthesis, and leaves (except of evergreens) in autumn ; 243. Decltii'ite (--i/iis). or I),, •Hi/nl. Bent or curved downward or forward. Decinit/inini'/. Several times compound- ed or divided; 10-2. 104. !>, composites. I decompound. /i, ,',1,11/11 nt (-ens). Reclining, but with summit ascending; 53. l>< ,-iii-rt nt (-nix), l>«-nrfirt. Running down into: as where leave- are seem- ingly prolonged below tbeir insertion, and so run down the stem. Decufsnte (-(ttux). In pairs alternately crossing at right angles. Dedvplication, Fr. Dedoublement. Same as Chorisis; 202. Dejinitt (-itus). Of a lixed number, not exceeding twenty ; or of a tixed or>ier. lit liitltt J tijturesctnce. Where axes of inilorescence end in a flower; 144, 151. Dt:jli-j:i:( or.-iiun. Downwards. Depauperate (-atus). Impoverished; as if starved; or diminutive for want of favorable surroundings. Depressed (-UK). Having the appear- ance or shape as if flattened from above. Derma. Greek for skin or surface of a p'ant or organ. DrKcmiliin/ t-ens). Tending or turning gradually downward. />< .«; nilin ,/ Axis. Primary root: 11. Detn -iniii'it, . Limited in number or ex- tent; as are the axe* of determinate inflorescence; 144, 151. I>,.-;'iit< UK. Terminating in. Desmos. Greek for things bound, or as if chained together. Dextrorse (l>,.rtri>rftiK: adv. Dextror- suni). Toward the right hand, or re- lating to it; 51. 140. />!. !>:.<. In Greek compounds, two, or double. Diachtnium. Synonym of Crsmocarp. GLOSSARY. 407 Diadelphia. A Linnaean (335) class having the sbimens. Diiult/phous (-us) Combined by their filaments into two sets; 250. Duii/iwsis. A brief distinguishing char- acter. DinypctttlcR, 341. Dialypetalous (-us). Same as polypeta- luiis, i. e. of separate petals; 244. Dialyphyllvus (-us). Bearing separate leaves. Diandrin. A Linnaean class with per- fect dowers having only two stamens; 334. Diandrous (Diander, £c.). Having two stamens; 249. Diaphanous (-us). Letting the light shine through. Dicarpellary. Composed of two carpels or pistil-leaves; 201 Dichusium. A two-parted or two-rayed cyme; 152, 155. Dichlamydeous (-eus). Having a double perianth; 191. Dichotomous (-us). Forked in pairs; two-forked. Dichdyamous (-us), Dichogamy. Her- maphrodite with one sex earlier de- veloped than the other in the blossom; 219. Diclesium. Name of a fruit consisting of an achenium within a separate and free covering made of perianth, as that of Mir.ibilis. Diclinous (Diclinis). When flowers are of separate sexes ; 191. Dicoccous (-us). Fruits of two cocci. Dicotyledons, Dicotyledones. Plants of the class marked by having two coty- ledons ; 27, 339. 340, 344. Dicotyledonous (-eus). Having a pair of cotyledons; 10, 314. Didyrnous (-us). Twin, found in pairs. Didynamin. The Linnaean class marked by didynamij (335), i. e. Didynamous (-us). When a 4-androus flower has the stamens in two pairs, and one pair shorter than the other; 250. Diercsilis. Mirbel's name for a dry fruit composed of several cells or car- pels connate around a central axis, and separating at maturity, as that of Mallow. Difformis. Of unusual formation. Diffuse (-usus). Widely or loosely spreading. Diyamous (-us). Of two sexes in the same cluster. Digitate (-atus). Fingered ; a compound leaf in which all the leaflets are borne on the apex of the petiole; 101. Ditjitctttly. In a digitate mode; same as Palmately. Digitate-Pinnate, 104. Ditjijnia. A Linnaean order character- ized by having the gynoecium Diijijnous. With two separate styles or carpels; 2G1. Dimerous (-us) Of two members in each circle; 176. Dimidiate (-atus). Halved, or as if one- half was wanting. Dimorphous (-us), Dimorphic, Dimor- phism. Occurring under two forms; 225, 234 Dicecia. Linna?an class (355) of plants with the flowers Diuecious (Diascius, Dioicous). Unisex- ual, and the two sexes borne by dis- tinct individuals; 191. Diwcio-pulyyamous. When some indi- viduals baar unisexual and others bi- sexual flowers. Dipttalous (-us). Two-petaled; 244. Diphyllvus (-us). Two-leaved; 243. Dlpln. See Duplo. Diplostemonous, Diplostemoni/. Having twice as many stamens as petals or sepals ; 177, 198. Diploteyium. A capsule or other dry fruit, invested with adnate calyx; an inferior capsule. Dipterous -(us). Two-winged. Dire nipt ion (-io). Syn. of Chorisis; 202. DisciJ'erous (-us). Disk-bearing. Discijbrm (-onnis). Depressed and cir- cular, like a disk or quoit. Discoidal or Discoid (Discoideus). Ap- pertaining to a disk. A discoid head is one destitute of ray-flowers. Disc or Disk (Discus). A word used in several senses. The disk or c'Jjc of a flower is a development of the torus within the calyx, or within the corolla and stamens ; 213. In a capitulum or head of flowers it is the central part of the cluster, or the whole of it as opposed to a border or ray. It is the face or surface of any organ, such as a leaf-blade, as opposed to the mar- gin. In vegetable anatomy, certain round spots or markings on cell-walls are termed discs. Discolor. When the two faces of a leaf, &c.. are unlike in color. Discrete (-etus). Separate ; not coales- ced. 408 GLOSSARY. Disepalous (-us). Two-sepaled ; 244. Disk-jlowers. Those belonging to the di~k, or body, and not to the margin or ray of a eapitulmn. Dissected (-us). Deeply cut or divided into numerous segment-. Dissepiment (-tntum). A partition in an ovary or pericarp; 264. Dissilii-nt (-ens). Bursting asunder or in pieces. Distil-In, it.-: (-us). Disposed in two ver- . tical ranks ; 122. Distinct ( Distlnctus). Separate from ; not united. Distractilt (-His). Carried widely apart. Diilucous (-«.-•)• Of two thecae, or cells, as are most anthers; 254. Diurnal. Daily ; occurring in the day; sometimes used for ephemeral. Diraricate (-ntus). Extremely divergent. Divergent, Diverging (-ens). Inclining away from each other. Divided (Divisus). Where lobing or segmentation extends to the base; 98. Dodeca. Greek for twelve. Used in Dodecaijynia. Linnoean order with flow- ers. Dudeciiyynous. Having twelve styles or distinct carpels. Dodecdmerous (-us). Of twelve parts in the circle. Dodecitndria. A Linnxan class (334) with the flowers. Dodecdndrvus, Having twelve (or from 12 to 111) stamens; 249. Dodrantulis. A span (about nine inches) long. Dolnbriform (-ormis). Axe-shaped or hatchet-shaped. Dorsal (-nils). Relating to the dorsum or back. Dorsal Suture. That which answers to the midrib of a carpel ; 201. Dofsij'i runs. Borne on the back. Double. Has a technical use when a flower is said to be "double;" this denoting one in which the leaves of the tlower are monstrously increased mostly at the expense of the essential organs. Downy. Pubescent with fine :ind soft hairs. LOOM-IN- synonymous with soft- pubescent, tomentoxc, iV-c. Drepdnium, .\ su'kle--.|iaped cyme; 156. Driipii<-i mis (-i'iis). Resembling or relat- ing to a drupe. Drupt- (l)rupn). .\ stone fruit; 297. Drupelet, Drupel (Drupvola). A dimin- utive drupe; 297. Drupetum. An aggregation of drupes; 300. Duct. In vegetable anatomy, an elon- gated cell or tubular vessel, lound espe- cially in the woody (tibro-vascular) parts of plants. Dumetuse (-vsus). Pertaining to Dume- tum, a thicket, or Dumus, a bush. Dumose (-usus). Bushy, or relating to bushes. Duplo. Twice as many, in Greek com- pounds, Dijil". Duramen. The heart-wood of an exo- genous stem ; 80. Dwarf. Of small size or height com- pared with its relatives. Dyclesium. See Diclesiuin. E- or Ex- As a prefix to Latin words, carries a privative meaning, as Eco&- tate, without ribs, Exalbuminous, with- out albumen. Eared. Same as Auriculate. Ebracteate, Ebracteolate (-atus). Desti- tute of bracts or bractlets. Eburneus. Ivory-white. Ecdlcarate (-atus). Spurless. Echinate (-atus). Beset with prickles, like a hedgehog. Echinulate (-atus). Beset with diminu- tive prickles. Edentate (-atus). Toothless. Effete (-etus), or EJfaetus. Past bearing; fund ionless from age. Efflorescence (-entia). The time or state of blossoming; anthesis. l-'.J'nse (-usus). Very loosely spreading, more so than diffuse. Eglandulose (-osus). Destitute of glands. I-:is). In the form of an el- lipse. Oval or oblong with regularly rounded ends ; 95. Emari'i'lus. Flaccid or withered. Enwryinate (-ntns). With a notch cut out of the margin; or, as usually ap- plied, out of the extremity; 97. Embracing. Clasping by the base. GLOSSARY. 409 Embryo or Embryon. The rudimentary plautlet formed in a seed ; 9, 311. Embryonal. Relating to the embryo; as Embryonal Vesicle ; 284. Embryo-sac. The cell in the ovule in winch the embryo is formed; 283. Embryuyeny. Embryo-formation. Emersed (Emersus). Raised above and out of the water. Enantiubldstus. With embryo at the end of the seed diametrically opposite the hilum. Enation. Having outgrowths from the surface, Sac,. ; 179. Endeca. In Greek compounds, eleven; as in Endecdndrous, Endecdgynous. With eleven stamens or eleven stvles, &c. Endemic. Confined geographically to the particular region. Endocarp (-drpiurn). The inner layer of r a pericarp; 288. Endochrome (-oma). Peculiar coloring matter in cells; especially the color- t ing matter of Algae. Endoyens, Endoyence. Endogenous Plants; 70. Endoyenous structure, 70. Endopleura. Inner seed-coat; 306. Endophlmim. Inner bark; 77. Endorhizal (-us). Said of an embryo which has the radicle sheathed by the cotyledon or plumule wrapped around it in many Monocotyledons ; hence Endorhizce. Synonym of Monocotyle- , dones. Endosperm (-ermium). Synonym of the albumen of a seed ; or the inner albu- / men ; 14, 310. Endostome (-oma'). The foramen of the inner coat of an ovule ; 277. Endothecium. Inner lining of the cell of an anther. Enervis, Enervius. Nerveless; no ribs or veins visible. Ennea. In Greek compounds, nine ; as in Ennenrjynia. A Linnjean ordinal name, and Ennedr/ynnus. With nine separate styles or carpels ; 337. Ennenndrin. Linnaean class, and Enne- nnr/rous, with nine stamens; 249, 334. Enodnl (Enodis). Without a node. Ensdtus. Same as Ensiform. Ensiform (-ormis). Sword-shaped ; i. e. like a broad sword, or the leaf of an Iris. Entire. Without toothing or division; the margin whole and even ; 97. Entomophilous. Said of flowers which are habitually fecundated by pollen t carried by insects; 217, 218. Entopltytes (Entophyta). Plants grow- ing in or out of other plants, as cer- tain Fungi, &c. ; 4. Ephemeral. Lasting only for one day. Epi. In Greek compounds, upon. Epiblast (-us). Name sometimes given to the first (and an undeveloping) leaf of the plumule of the embryo of grasses and grain. Epiblastema. A superficial outgrowth from leaves, &c. ; 21 0. Epicalyx. Name sometimes given to an involucel resembling an accessory r calyx. Epicarp (Epicarpium). The external layer of a pericarp ; 288. Epichilium. The terminal portion of the labellum of an Orchid, when this is of two parts. Epidinal (-us). Upon a torus. Epicdrolline. Upon a corolla. Epidermis. The skin of a plant; 76, 89. Epiyceous (-eus). Growing on or out of the ground. Epigynous (-us). Literally on the pistil; meaning on the ovary, or seemingly so; 183. Epipetalous (-us). Borne on (adnate to) the petals ; also used in the sense of placed before the petals. Epiphlceum. The outermost or corky bark; 76. Epiphyllous (-us). Growing on leaves. Epiphytal. Pertaining to Epiphytes (Epiphyta). Plants growing on other plants by way of attachment, but not parasitic; air-plants; 35. Epipterous (-us). Winged or wing- bearing at summit. Episepnlous. On the sepals; also used in the sense of standing before a , sepal. Episperm (Epispermium). The coat or , outer coat of a seed ; 305 Epitropovs (-us). Name (by Agardh) of an anatropous ovule with rhaphe averse when ascending, adverse when suspended; 282. Equal (dSqualis). Alike as to length of , number, &c., as the case may be. Equitant (-ans). Riding; folded around, as if straddling over; 108, 138. Erect (Erectus). Standing upright, mostly in relation to the ground. 410 GLOSSARY. sometimes when perpendicular to the surface of attachment. l-'.ritin. Greek for wool; used in com- pouml words, as I-'. 1 1 ni ill us. With woolly flowers. Eridphorus. Wool -bearing. l us. Woolly-leaved. ( AVo.v//-/.*). l!eak!e-<. roiuisus). Covered with a meal- like powder. Fttsciate (-atus). Said of monstrous ex- pansions of stems, giving the appear- ance as of several stems coalescent in one plane. Fusdr/i (-if/tin ). A close cluster or bundle, whether of tlowers, stalks, roots, or leaves: 147, 153. Fiisi-'n-li it ( Fasciculatits, Fascicularis). In a fascicle; 131. Fed : same as Alveolate. Feather-veined. Having veins all pro- ceeding from the sides of a midrib. Feathery. See Plumose. GLOSSARY. 411 Fecula or Fcecula. Starch-like matter. Applied to a pistillate flower, or to a pi. nit producing only such flowers. Fenestrate (-atus), Fenestralis. Pierced with large holes, like windows. Ferruyineous or Ferruginous (Ferru- gincits). Colored to imitate iron-rust. Fertile (-His). Fruitful, fruiting, or ca- pable of producing fruit; as a fertile flower is one provided with a well- formed pistil ; 191. In English descrip- tions, Flower fertile usually means a pistillate or female flower. Stamens or anthers are also said to be fertile when polliniferous and capable of fer- tilizing. Fertilization. Synonym of fecundation, as of the ovule by pollen; 215. Fibre (Fibrn). Any line filament; Ihe elementary components of wood, £c. ; delicate roots, &c. Fibril (-ilia). A diminutive fibre. Fibrillnte (-atus), Fibrillose (-osus). Furnished or abounding with fibres or fibrils. Fibrous, Fibrose (-osus). Composed or of the nature of fibres. Fibro-i'itscular. Consisting of woody fibres and ducts. Fiddle-shaped. Obovate and with a sinus or contraction on each side. Fidus. A Latin termination for cleft or lobed. Filament (-entum). The stalk or sup- port of an anther; 165, 251. Also any fibre-shaped or thread-like body. Filamentous, Filnmentose (-osus). Com- posed of threads or filaments. Filicoloi/y. The botany of Ferns : re- placed by Pteridology. Filiform (-ormis) . Thread-shaped; long, slender, and terete. FiUpendulous (-us). Hanging from a thread. Fimbria. A fringe, or dissected border. Fimbrinle (-atus). Fringed ; bordered by slender processes or marginal ap- pendages. Fimbnllule (-atus), Fimbrilliferous (-us). Bearing Fimbrillce or diminutive fringe. Fingered. See Digitate. Fissiparous. Multiplying by the divi- sion of one body into two, and so on. Fissus. Split or cleft. See Fidus. Fistular, Fis/ulose (-osus). Hollow through the whole length, as the leaf and stem of an Onion. Flubellate (-aius), Flabelliform (-ormis). Fan-shaped ; much dilated from a .wedge-shaped base, and the broader end rounded. Flabellinerved. With radiating straight nerves ; 92. Flagellate (-atus), Flayellaris. Produc- ing filiform runners (Flayella), or runner-like branches. Flagelliform (-ormis). Runner-like; long, slender, and supple like a whip- lash or Flagellum ; 53. Flammtus. Flame-colored. Flavescent (-ens). Yellowish or pale yellow. Flitvus. Pale yellow or ochre-yellow. Fleshy. Succulent ; of the consistence of flesh. Flexuous, Flexuose (-osus). Zigzag: bent alternately in opposite direc- tions. Floating. Borne on the surface of water. Floccoxe (-osus). Bearing or clothed with locks of soft hairs or wool (flocci). Flocculent. Diminutive of floccose. Flora (Goddess of flowers). The aggre- gate of the plants of a country or dis- trict; or the name of a work which systematically describes them; 369. Florid. Belonging to the flower. Floral Envelopes. Floiver-leaves; 164. Floret. A small flower, one of a cluster Floribundus. Abundantly floriferous. Floriferaus(Florifer, Floriferus). Bear- ing flowers Florida. A small Flora; the Flora of a restricted district. Flos. Latin for flower. — Flos plenns. A " double " flower; that is, one in which petals are increased abnormally, com- monly at the expense of the androe- cium or the gynoecium also; 171. Flosculus. Latin for floret. Flower. The whole reproductive appa- ratus in a phsenogamnus plant ; 163. Flower-bin}. An miexpanded blossom or undeveloped cluster; 40. Flowering Plmls, 3, 344. Floiverless Phnts, 3, 344. /•'In/ Inns. Floating. Fluriiit/le, Fluriiitilis. Belonging to a river or running water. Fly-traps, 113. Fcemineus. Feminine or female flower, plant, &c. ; 191. Foliaceons (-etis). Leaf-like in texture or appearance; or bearing leaves. Fu/i'ir (Fo/inris). Relating to leaves. Foliation (Foliatio). Leafing out. Foliate (-atus). Having leaves. With Latin numerical prefix, bifoliate, iiii"x/>i ring, Gymnospermce. A sub- class iif naked-seeded plants; 268, 344. Gytniiini/ii minus (-11*). Naked-seeded, as opposed in Angiospermous. fii/iiiiin/nii. .\ Linnaean class, character- ized by the (lower In in- Gynandrous. Stamen- borne mi ludnate to) the pistil, even to the style or ma; 251, 335. Gynabnse ( Gynobasis). An enlargement or prod uei ion of the torus on which the gyiufciuin rests or is somewhat elevated; 212. Gyno-di&cious. Diitcious with some flowers hermaphrodite and others pis- lillate only; 191. Gynacium. The pistil or collective pis- lils of a (lower; the female portion of a (lower as a whole; 165. Gynophore (Gynopliornm). The stipe of a pistil ; 212. fii/iin.-i/i i/iiini. A sheath or covering of the gyiiiccium, of whatever nature. GyniiKti in in in. The column of an Orchid, consisting of androecium and summit of the g\ meciuin combined. Gyriiti- (-nt UK). Curved into a circle, or taking a circular course. Gyrn.-n (-uxim). Curved backward and forward in turns. (ll,ibitiiK). The general appear- ance of a plant. . Habitation; the geographical limits or station ; .'iiiii. Hii'inntltii: (-/<•;/.<). Brown-red. II! ii- or ll'iUn i -i /-.i hi/us (-us). With only one row of pe'aU. Haplostemonovs (-us). With a single -cries of stamens; 177. J/ii*:ntr (-ntus), f/astilis. Halberd- shaped, like the head of a halberd, i. e. sagittate, but the basal lobes di- rected outward or at right angles to the midrib of the leaf; 90. /// nd. The form of inflorescence termed <'"j'ituln/it, viz. a cluster of sc^-ilc flowers on a very short axis and centri- petal in evolution; 147. ll< urt-thaped. Ovate with a sinus at base; 96. Heart -wood. The older and matured wood of an exogenous stem ; 80. Hibetntf (-ntus). Having a dull or blunt and soft point. JJtlicutd (-oidi «.ous (-us). Same as amphitro. pous or half anatropous. Hijitit. The (.ireek numeral seven. Heptayynia. A Linnaean artificial order, having seven stj-les or distinct car- pels; 337. Hijitii/itcrous (-us). Of seven members. Hi jitnnilrin. The Linnrcan class with seven stamens ; 334. Heptandrous. Seven-stamened; 249. Herb (Herba). A plant with no persist- ent woody stem above ground; 50. Herbaceous. Of the texture, color, or other characters of an herb. //( r/tiiriinn. Hi rbnl. A collection of dried specimens of plants, systemati- cally arranged; 380. Hi ri-i'ii/n minis (-us). Said of hermaphro- dite flowers when some structural ob- stacle prevents autogamy. Hi rum/ill roilite (-it us). Of both sexes; 191. //i.yit riifinm. A hard-rinded berry, like an orange and lemon ; 21)9. lltiiracmy. Svnonvm of Dichogamy; 219 Hi ti rns. In Greek compounds, denotes diverse or various, as Hi f, rnrin-fiinis (-us, IJeterocnrpicus). Producing more than one kind of fruit. GLOSSARY. 415 Heterocephalous (-us). Bearing two kinds of head or capitulum. Heterodite (-itus). Anomalous in forma- tion. Httt roc line (-inus). Nearly same as Heterocephalous ; on separate recepta- cles. Htterodromous (-us). Spirals of changing direction. Heteroyamous (-us). Bearing two kinds of flowers. Heteroyrneous. Not uniform in kind. Heteroyime or Heteroyoneus. When the flowers are dimorphous or trimor- plunts as respects relative length, &c., of stamens and pistil ; 225, 234. HeterostyU-d. Same as Heterogone ; 234. Heteromerous. Of members not corre- sponding in number. Heterophyllous (-us). Having leaves of more than one form. Heterotropous (-us). Turned in more than one direction, or in an unusual di- rection; same as Amphitrdpous ; 279. (Also used by Agardh for collateral ovules tinned back to back; 282.) dexa. Greek numeral six ; from which is formed tiextujynia. Linna?an artificial order, of flowers with six styles or distinct car- pels ; 337. Htxdyynous. Having the character of Hexagynia. Hexdmerous (-us). Of six members; 176. Hexandria. Linnaean class with perfect flowers of six stamens; 334. Hexdndrous. Having six stamens ; 249. Hexapetalous (-us). Having six petals. Hexapliyllous. Six-leaved. Hi j-i'ipterous (-us) Six-winged. Hexascpalous. Having six sepals. Hexaslt'monous. Having six stamens. Hibernaculurn. A winter-bud; 40. Hidden. Concealed from sight ; as Hidden-reined, where the veins are in- visible, as in the leaves of Pinks and Houseleeks. Hii-miil (-nils). Relating to winter flilttr ( Hilaris). Belonging to the hilum. Hilum. The scar or place of attachment of the seed ; 277, 305. Hippocrepiform (-ormis). Horseshoe- shaped. Hirsute (-utus). Pubescent with rather coarse or stiff hairs. Hirtellous (-us). Minutely hirsute. Hirtus. Hairy, nearly same as Hirsute. Hispid (-idus). Beset with rigid or bnstJv hairs or with bristles. Hitpidulous (-us). Minutely hispid. Hoary. Grayi.'-h-white with a fine and close pubescence. See Canescent. Hoiosericeous (-ens). Covered with fine and silky pubescence. Homocarpous (-us). With fruit all of one kind. Hoinddromous(-us), Humodromy. With spirals all of uniform direction. Homoyamous (-us). Bearing one kind of flowers. Homoyeneous. All of one nature or kind. Homoyonous or Homoydne. Homomor- phous as respects the stamens and pis- til; opposed to dimorphous; 225. Homoloyue. A homologous organ or part. Homologous. Of one name or type, such as leaves and parts answering morpho- logically to leaves ; 6. Homomdllus. Said of leaves and the like which are all turned in one direction. Homomorphous (-us). All of one form. Homostyled. Same as Homogone. Homdtropous (-us). Curved or turned in one direction ; applied also to the ein- brvo of an anatropuus seed, with rad- icle next the hilum ; 312. Hood. See Cucullus. Hooded. Bearing or in form of a hood. Hoi-nus, Horndtinus. Of the present year. Horny. Of the consistence of horn. See Corneus. Hortensis, Hortuldnus. Pertaining to the garden. Hortus siccus. Old name of an herbarium. II ami. On the ground. Humifusus, Humistrdtus. Spread over the surface of the ground. Humilis. Low of stature. Hijnline (-inus). Transparent or trans- lucent. Hybrid. A mongrel, or cross-breed of two species; 321. Hydrophytes (Hydrophyta). Water- plants. Hyemdlis. See Hiemalis. Hypanthium. An enlargement or other development of the torus under the calyx: 214. Hypo. In Greek compounds, denotes under, beneath, lower. Hyptinthodium. Same as Syconium; 149, 303. Hypticlitiium. The basal portion of the labellum of an Orchid. Hypocraterimorplious (-us), or IJypocra- teriform, but the latter is a hybrid of 416 GLOSSARY. Greek and Latin. Salvcrform or sal- ver-shaped ; that is, in the form of a saker rai>ed on a central support or -ii-in beneath. Said of a corolla and the like wiih slender lulu- abruptly ex- into a Hat linib; 248. * (-II'IIK). Growing or remain- ing iiiidei-gTuimd ; 111. //.'//<"//.'/ '""'•< (-«.<). 1'iuler or free from the g\ iHreinm or pi-til ; 182. HypopiiyUous (-US). Growing -on t lit- un- der .-ide of a leaf. Hypopky Ilium. An abortive leaf or scale under another 1. al. nr -ccniing leaf, as in Asparagus and Kuscus. Hypsophylla. Answers to the German " Ilochbliitter," or high leaves, those of the inflorescence, t. t- bracts and the like; (i. Hysteranthous (-«.<). With leaves pro- duced later than the blossoms. Icosandria. The Linna?an class with twenty stamens (as the name denote- 1 or a larger number, inserted on the calyx; '.r.j4. /cosandrows is the corre- sponding adjective; 249. /lulu rliix. Not bearded. Imbricate (-utus), Imbricntivc. Over- lapping so as to '' break joints," like tiles o'- shingles dii a roof; either with parts all in one hori/onta! row or cir- cle, as in the a-stivation of a calyx or corolla, when at least one piece must be wholly external and one internal; or with the tips nt lower parts covering the bases of higher ones in a succession of rows or spiral ranks; 135. Iniin,iri,intil,' (-ntiia). Not margined or bordered. tin in i r.-ti-tl (-us). Growing wholly under water. /HI/HI ri-/>in»iili'. Pinnate with an odd terminal leaflet : 101. liiii'i/iiiln/i-rtil (-H/is). Unequal-sided. /ii'iiiix. Fnipty, as an anther containing no pollen. fnappendiculate (-ntii.t). Not appen- daged. In,- in, f,-i ni. Same as Canescent. Hoary-white. (-uliif). Flesh-colored. See Cameus. Inrisi'il (-its). Cut irregularly and sharply; 98. Included (Inclusus). When the part in question does not protrude beyond the surrounding organ. Incomplete (-us). Wanting some essen- tial component part : 190. /Ki-rnt.'ii/i '->'///.<). Thickened. Ini-nliS;c. /«/'( rim- ( I nft rii.-:). Said of ono organ when below another. In the blossom also in the sense of anterior ; 100. An inferior calyx is one below the ovary, or free; 18't. An inferior ovary is one with adnate or superior calyx ; 183. /H/tn/x/ (-ntii.t). Hladdery. lull, .1; ,1 (-».•;). Bent or turned abruptly inward. Ii(tli>r<.*rinn'. Mode of di-position of ildwers ; less properly used fbraflower- . cluster itself; 141. Fnfroi-amllary (-arts). I?elow the axil. [nfvndibuliform (-ortnts), Infundibular (-aris). Funnellbrm, funnel-shaped; 249. GLOSSARY. 417 Innate (-atus). Borne on the apex of the supporting part; in an anther the counterpart of adnate ; 252. Innovation (-io). A new-formed shoot. Inosculating. Same as Anastomosing. Inseparate, Inseparation. Terms pro- posed by Masters to express coales- cence; 181. Inserted (-us). Attached to or growing out of. Insertion (-io). Is the mode or place where one body is attached to that which bears it. Integer. Entire in the sense of un- divided, or not lobed ; 97. Iittvgi'rnmus. Entire in the sense of quite entire, i. e. the margin without dentation; 97. Inter. Between; as in Intercellular, between the cells, &c. Interfoliaceous (-eus). Between the leaves of a pair, as the stipules of many Rubiacese. Internode (-odium). The portion of stem between two nodes; 6. Interpetiolar (-aris). Between the pet- ioles. Interruptedly pinnate. Pinnate with- out a terminal leaflet. Intine. The inner coat of a grain of pollen. Intrafuliaceous (-eus). Within or be- fore a leaf. Introfiexed (-us). Same as inflexed. Introrse (-orsus). Turned inward or toward the axis ; 253. Intnn-enius. Same as hidden-veined. Jntruse (-usus). Pushed or projecting inward. Involucellate (-atus). Provided with a secondary involucre or Involucel (-illlum). An inner or secon- dary involucre that of an umbellet, &c."; 142. Involiicrate (-atus). Provided with an involucre. Innilucre (fnvolucrum). A circle of bracts subtending a flower-cluster; 142. Involute (-utus). Rolled inward; 133. Irregular (-nris). Exhibiting a want of symmetry in form; 184. Irregularity, 179, 184, 219. Isadelphous (-us). Equal brotherhood, as when the number of stamens in two phalanges is equal. Ifnchrous. All of one color or hue. jsomerous \-us'). The members of suc- cessive circles equal in number ; 175. Isostemonous (-us). The stamens just as many as the petals, £c. ; 177. Jsoslemony, 196. Jointed. See Articulated. Juba. A loose panicle, with axis de- liquescent. Juyum, p\.juya. A pair of leaflets. So pinnate leaves are unijugate, with a single pair of leaflets; bijuyatc, with two pairs; trijuyatt, with three pairs or juga, &c. Also the ridges on the fruit of Umbelliferae are termed juga. Julus. Same as Amentum or Catkin. Julaceous (-eus). Catkin-like, Amen- taceous. Keel. A central dorsal ridge, like the keel of a boat. The two anterior petals of a papilionaceous corolla, which are united into a body shaped like the keel or the prow of a vessel; 185. Keeled. Having a keel. See Carinate. Kernel. The nucleus of an ovule, or of a seed, i. e. the whole body within the coats. Kermesinus. Of the color of carmine. Key- fruit. See Samara; 294. Kidney-shaped. Crescentic with the ends rounded; very oblately cordate; 96. Kingdom, 325. Labellum. One of the petals of an Or- chideous flower, which is unlike the others. Labiate (-us). Lipped, mostly Bilabiate ; 247. Labiatiflorous (-us). Said of certain Composita; with bilabiate corollas. Labiose (-osus). Said of a polypetalous corolla which has the appearance of bilabiation. Labium. See Lip. Ldi'i'rnte (Lncerus). Irregular cleft as if torn or lacerated. Lacinia. A slash; used for a slender lobe. Laciniate (-atus). Slashed; cut into narrow incisions. Lacinula. A diminutive lacinia or nar- row lobe. Lactescent (-ens). Yielding milky juice. Lactvus. Milk-white. Ldcunose (-osus). Abounding in pits, holes, or depressions (lacunae). Lacustrine ( Locust ris). Belonging to or living in lakes or ponds. 418 GLOSSARY. /." riyate (-atus). Smooth as if polished. Lcevis (this form, and mil /• <•/.<, h.-is al- ways been used in botany). Smooth in tin- sense of not rougli. Lageniform (-ormis). Shaped like a 1-loreiice Hask or a gourd (the fruit of Lagenaria). Layouts. Hare-footed. Densely cov- ered with long hairs. Lnut l/ii. A thin ]>late. Ldmellur (-aris), LminUate (-atus), La- mtlluse (-osus). Composed of thin plates or lamellae. Lamina. The blade or expanded part of a leaf, £c ; 85, 245. Lunate (-atus), Lanuse (-osus). Bearing long and implexed hairs or wool (lana ). Lanceolate (-atus). Shaped like a lame or spear-head; narrower than oblong, and tapering to each end, or at least to tlie apex; 95. Laniit.innii* (-<>sus). Cottony or woolly ; dollied with soft and implexed hairs or down (lanue/o). Like a bur (lappa). Woolly-flowered. Latt nt. Undeveloped or dormant, as certain buds ; 40. Lateral (-fills). Belonging to or borne on the sides. Lateritious (-ius). Of a brick-red color. Latex. Proper juice, milky juice, and the like. Latirij'i-rous. Containing or conveying latex. L'ltitii'jitus. With broad partition. Lavender-color. Pale blue with some gray. Lams 1/iose. Leaf. The principal sort of appendage or lateral organ borne by the stem or axis : 85. lA-iif-blailc. The lamina of a leaf. l.i "l'-linii/iitt (-otus). Beset with small scurfy scales. J.ijiis. Greek term for a scale. Lfj>tos. Greek for slender; as in Lep- ti jilnjlliif, slendiT-leaved. Ltucvs. Greek for white ; whence Leuc/mt/ius. White-flowered. Leucophyllus. White-leaved, &c. Liber. The inner and often fibrous bark ; 77, 81. L'ul. See Operculum. Liijneous (-cus), Lujnosus. Woody. Lit/uli (/.ii.u/'i }. \ strap or stra]>- shaped liody. such the princi])al part of a ray corolla in Composite. The thin and scarious projei'tion from the summit of ihe sheath of the leaf of Glasses, ^c. : lOU. Or a similar out- growth of the inner face of certain petals; -Jll. Liyulate (-atus), Liguliform (-ormis). l-'uriiishi'd with a lignle; 148, 247. Ligutiflorout (-us). Said of the head of those Composite' which contain only ligulate corol'as. J.i/im-i mi.-!. Lily-like; 246. Limb (Lhnlins). A border, i. e. the ex- panded part of a gamophyllous peri- anth, &c., as distinguished from the tube and throat : 245. Sometimes the term is applied to the lamina or blade of a petal or a leaf. Limbate (-atus). Bordered. Line (LiiH'ii). The twelfth part of an inch. By some reduced to the tenth of an inch ; but the decimal line is un- usual in botanical measurement. Lhn'ir (-ri;-/.en>. l/. rii-iirji ( -injiiuin). One of the akene- like carpel> or a clo.-ed lialf-t'ruil ot' I'mbellifera' ; 2117. Mi-i-iiiiiuitif. Dividing into parts or sim- ilar portion*. I/, riilni/ltiK. Synonym of Internode. Mi /•<,«.< (-us). In Greek compounds, de- notes part* or members; hence Dime- runs. of two parts, £c. Mi.««-nr/i (-in-jiium). The middle layer of a pericarp; 285. Mesophlceum. The middle or green bark ; 76. Metamorphosis, Metamorpky, 167. Mii-rii/iil/f (-///")• 'I1'11' sl"'t or point in tlie >eed at which was the oriticeof the ovule; 277, 305. Mii-i-iiKjiiH-, . The smaller kind of spore in Lycopodiaceae, £c. Mill rili. The central or main rib; 92. Miiiiutt- (-n//if). Vermilion-color. Mi*/ a* or Mi.ii/iK. A cross-breed ; 321. Mitrii'/'iiriii or .^fi/rifunii (-OTmis). Mi- tre-sha]ieil or cap--haped. M^/nii/il/iltiii. The I.inii:eaii class con- taining (lowers with ftfonadelphotts stamens, /. e. those united by their lilaments into a tube or column ; 250, 335. Mni/itiii/riii. The l.inn:ean class (334) coiilainiiig (lowers with Mmim/i/rous, that is, a single stamen; 24!). Miillil/l//ii'll.-< ( -//M. ( )|le-tlo\\ere(l. Mi 111 Hi /'iirin (-a r 111 IK). Necklace-shaped; cylindrical and with contractions at intervals. \fonocarpellary. ()f one carpel; 201. Sfonocdrpic (-/(•//.<). Hfonocarpovs} .!/"•'»- iii-/ir/ii'ii>. ( >nly once fruiting: .".:!. Minim-! /iliiili>ii.< (-U8). Hearing a single capitulum. A cyme with one main \fonochlamydeous (-. fotuecitiiif ',-'«.-•'. or Monuicous (-«*'). With stamen> and pi>tils in separate Moroni- on the smie jilant : l!'l. Name of a Linn;ean arti- licial order, in class Syngeiie-ia; 337. Monograph. A -\-t'-matK account of particular genus, order, or other group; 369. Monoyynia. Name of a Linnaean arti- licial order, with solitary pistil, or style; 337; hence, adjectively. M"i/<>- tjynous; 267. Mninmn rtyle. Monosymmetrical. That which can be bisecled into equal halves in only one plane; 175. .l/..;/r;/.iri>».s- (-».<). Bearinu' progeny (frniiiiiLri only once, as annuals and biennials : 33. M»n*/t r ( Mi'iif/runi). A monstrosity, or unnatural development. Morphology, -ri- M»Ki-liiit< (-Hi/if). Exhaling the odor of musk. Mni-itiit/liintiK. Mucilaginosus. Slimy ; of (lie consistence or appearance of mucilage. M/icrn, MiKTniiii/inii. A short and small tip to a leaf. &c. (-K). Many-headed; many shoots or stems from the crown of one root. GLOSSARY. 421 Multifarious (-us}. Many-ranked, as leaves in several vertical ranks. Multijid (-idus). Cleft into many lobes or segments. Multiflorous (-us). Many-flowered. Multijuyate (Multijuyus). In many pairs or juga. Multilocular (-aris). Many-celled or several-celled. Multiparrous. Many-bearing ; said of a several-branched cyme ; 152, 155. Multiple fruits. The fructification of a flower-el uster when confluent into one mass ; -301. Multiplication. Same as Augmentation ; lit), 200. Multiseriul (-alts), Multiseriate (-atus). In several series. Muricate (-at us). Hough with short and firm excrescences. J//n-Hv. Nutlet. A diminutive Nut. See Nucule. Nutitnt (-ans). See Nodding. Ob. Over against ; as a prefix denotes inversely or oppositely, as Obeompressed (-«.•>•). Flattened the other way, autero-posteriorly instead of lat- erally. Obctmical (-icus). Conical, but attached at the apex. Obcordate (-(tins). Inverted heart- shaped, the notch at the apex; 97. Obdiploslemonous (-us), Obdiplustemony. When the stamens are double the number of the petals, but the outer series opposite the latter; 198. Obimbricate. Imbricated or successively overlapping downward. Obldnceolate. Lanceolate but tapering toward the base more than toward the apex ; 95. trbliqiie ( OUiquus). Unequal-sided or slanting. Ollimij ( Oblonyus). Considerably longer than broad and with nearly parallel -ides; 95. Obi'irutt (-atiis). Ovate with the broader end toward the apex; 95. Obowid Solid ohovate. Obtec/n*. Covered by something. Obtn*i (-nun*.). Blunt or rounded at the extremity ; 97. Obtn*iii*<-nlii*. Somewhat obtuse ; dimin- utive of obtuse. Obvallattu. Guarded on all sides or sur- rounded MS it' walled in. O/ii-i r*< .(I/in I-KI 1 1/. Same as the prellxOb. Obr <>i-i//ii/, t-ii/ii*). With a circular patch or a ring of eolor. Ochraceous (-eus). Ochre color; light yellow with a tinge of In-own. Or/inn. Ocrea. A leg".in--haped or tul>iilar stipule or rather combined pair of -li)inle- : l"ii. Orl/n nit . Ocrnit, (-utii.t). Furnished with oclirea or sheaths: IIH;. Ochroleucotts (-».<). Yellowish-white or between white and yellow. Oct<>. F.ight. [n composition gives such term- as the following. ()<-/:i,:i/ii!ii. I.imci-an artilieial order with s( eight-sty led) Mowers ; 337. Octdmerous. Composed eight parts in the circle. Octandi-ia. Tlie Linmvan class with Oftmi'lrous, i. e. eight-etamened flow- ers; •J4'.». :;:J4. Octdni. In eights. Octopetaluus (-us). Eight-petalled. ( I, /l/iciiin/ (-nils). Used in medicine or the arts, therefore in the shops. Offset. A short lateral shoot for propa- gation ; 53. OUits, Oideus, -odes and -ides. Greek for likeness, used as terminations, in- dicate similarity to; as JJinntlioides, resembling a Pink. Oleraceous (H us). Esculent in the way _of a pot-herb. Olir/os. Greek for few; in compounds giving such terms as Oligdndrous (-us). With few stamens. Ollyanthoim (-us). Few-flowered. Oligdmerous (-us). Of few members. Oliyospermous (-us). Few-seeded. OUnii-i n* (-eus). Olive-green. Omphatodium. A mark (navel) on the hilum of a seed, through which passed vessels to the chalaza or rhaphe. One-sided. Either turned to one side, or with parts all turned one way, or unequal-sided. Oophoridium. The spore-case for the larger spores in Selaginella, &c. Opnque (O/nii-K.i). Mostly used in the sense of not shining or dull. t)/ti i-fitliiti (-H//I*). Furnished with an ( >perculum or lid. Opercvlum. A lid; a top which sepa- rates by a transverse line of separation as does that of a pyxis. Opposite (-ilus). Set against; as leaves over against each other when there are two on one node : or one part lie- fore another, as a stamen before a petal : C. ] •_>/i/in.i!/ij',itiii.i (-in.i). Plaeed <>]>|iosite a leaf, as is a tendril or peduncle in Vitis. ,\:c. Ojyx'titijit tuliius (-us). Placed before a petal. Oppositisepalous (-us). Situated before a -e]ial. Orbicular (-«m). Orfiii-iilnte- (-ntus). Said of a flat body with a circular out- line: 95. Orchidaceous, 246. GLOSSAEY. Order ( Ordo). Group between genus (or tribe) and class; 328. Ordinal. Relating to orders. Organogeny (Organogenesis). The for- mation or early development of or- gans ; 2. Oryanoyraphy, Oryanoloyy. The study of organs and their relations; 2. Organs of Vegetation, 11. On/ydlis. Six feet high, or of the height of a man. Ornithophilous. Said of flowers which are habitually fecundated by pollen brought by birds; 217. Orthoploceus. Said of an embryo when incumbent cotyledons are folded around the radicle, as in Mustard. Orthos, Greek for straight; whence com- pounds such as the following Orthostichies. Vertical ranks ; 121. OrtJiostichous. Straight-ranked. Ortkdtropous (-us), Orthotropal. De- notes an ovule or seed with straight axis, chalaza at the insertion, and ori- fice at the other end ; 277. Has been applied to an embryo with radicle pointing to the hilum; 312. Os (oris). The mouth or orifice. Osseous (-us). Of the texture of bone. Ossiculus. A little stone, same as Pyrena. Ostiolate (-atus). Furnished with a small orifice or little door (Ostiolum). Outgrowths, 209. Oml ( Oval-is). Broadly elliptical; 95. Ovary (Ovarium). The ovuliferous part of a pistil; 166. Ovate (Ovatus). Of the shape of the longitudinal section of a hen's egg, the broader end basal; 95. Used also for an egg-shaped solid. Ovoid ( Ovoideus). Used either for solid ovate or solid oval, more properly for the latter. Oculate (-atus), Ovuliferous. Bearing ovules. Ovule, Ovulum. The body in the flower which becomes a seed; 166, 276. Pagina. The surface of any flat body, such as a leaf. Palaceous (-eus). When the edges, as of a leaf, are decurrent on the support. Palate (Palatum). A projection in the throat of a personate gamopetalous corolla; 248. Palea. A chaff, or chaff-like bract, such as the chaffy scales on the receptacle of the head in many Composite ; also an inner bract or glume in Grasses; 142. Paleaceous (-eus). Chaffy ; furnished with paleifi; or chaff-like in texture. Paleola. A diminutive palea, or one of a secondary order ; one of the names of the Lodicule or Squamella in Grasses. Palculate (-atus). Furnished with pale- olse. Palets (Pales of some English botanists). Same as Palese; 142. Palmdris. A palm's breadth or length; i. e. equalling the breadth of the four fingers of the palm. Palmate (-atus). Lobed or divided so that the sinuses point to or reach the apex of the petiole or insertion; 101. Palmately (Palmatim or Palmati-). In the palmate manner. Ptilmately veined, 93. Palmatijid (-idus), Palmatilobate, Pcl- matistct). Palmately cleft, lobed, in- divided. Palmineroed. Palmately nerved ; 93. Pdludose (-osus), Palustrine (Palustrtf or Paluster). Inhabiting marshes. Pdndurate (-atus),Panduriform (-omits* See Fiddle-shaped. Panicle (Panicula). A loose compound flower-cluster, such as is produced by the branching of a raceme, or the ir- regular branching of a corymb ; 150. Panicled, Paniculate (-atus). In a pani- cled manner or borne in a panicle. Pannosus, Panniformis. Having the ap- pearance or texture of felt or woollen cloth. Papery, Papyraceous. Having the text- ure of paper. Papilionaceous (-eus). Butterfly-like; applied to a peculiar polypetalous corolla ; 184, 246. P « pillar (-aris), Papillose (-osus), Papil- late (-atus). Bearing or resembling papillae, minute nipple-shaped projec- tions. Pappiferous (-us), Pappose (-osus). Bearing a pappus. Pappus. Thistle-down; thence applied to various hairy tufts on akenes or fruits ; and thence to any production or structure which takes the place of the limb of the calyx on the akenes of Composite; 192, 295. Papuliferous (-;«), Papulose (-osut) Covered with Papulae, or small pirn- pies GLOSSARY. Unused name given to an abortive pistil ur carpel. J'iiriii;ii-nll,i. A crown or internal ap- pendage or deduplication of a corolla. I'n mill /-/tin-til, reined, &c. Same as Nerved; (J1. Parapetalous (-us). Said of stamens, \c., \\hicli stand at each side of a petal; 178, 201. }'u i-ujili i/sis, ]>\. 1'n nijih fists. Jointed thread-like bodies, ol no known tunc- tion, accompanying the archegonia of Mosses. / rasitic (-icus). Growing on or in and living uijon another plant or even ani- mal; '•><>. ]'iirn.-ti man. Name rarely applied to an abortive stamen or body in place of or accessory to a stamen; same as Sta- minodium. J'iirii."/n-/iitK. Secondary spirals in phyl- lutaxy; 127. Parenchyma. Common or soft cellular tissue. Parenchymatous. Of the nature of or composed of parenchyma. J'ur'n.t, pi. jiuritttm. The wall of any organ. J'liriiinl (J'm-ii tn/if). Borne on or re- lated to the wall; 265. J'liri/iiiitiiiti (-nt/m). Even-pinnate; same as abruptly pinnate; 101. I'lii-hil. 1'tirtitt (-itus). < 'left nearly but nut (juite to base; 98. Parthenogenesis, I'm-iln //<>i/< mj. Pro- duction of seed without the interven- tion of pollen; 285. 1'iiriinl ( /'iii-i/iilis). Secondary, as Par- tial involucre (142), peduncle (143), petiole (105), umbel (150). &i . ]'nr/i/>li (I'lirtiLilis). At length sepa- rating or easily to be separated. J'tirtlliun. In one sense a se]iarated por- tion or segment ; in another and the more usual, a wall or dissepiment. Patettiform \-orini*). Disk-shaped, cir- cular with a rim, of the form of the patella or kncepan. J'n/nii ( /'nit ii.i). Spreading; either widely open or diverging widely from an a \is. l'i//i i//i.-:tiiiiii.t. Superlative of Patens; extremely spreading. Pdtulous (-«>•) Slightly or moderately spreading. Pauciflorous (-//a). Pew-flowered. 1'iiUfijiiHus. Few-leaved. Pear-tliii/>< i/. < )ho\ oid or oliconical with more tapering base. Pectinate (-atus). Pinnatifid with nar- row and closely set segments, like comb-teeth. J '< 'lulls. A loot long or high. Pedutv (-utui). Pahnately divided or parli'd with the lateral divisions two- elett : re-embling a bird's loot. Pedatipartitus, -ln/mtiif. -Kn-i/i.-t, ^c. Pe- diitely parteil. lobed, divided, \, . J'"/ii;l (-iliiiti). An ultimate tlower- -talk or its division; the support of a single tlo\\er ; 143. /'< iln; II, iti (-utii.-t). IVdicelled, borne on a pedicel. I'liln-uliif. Name- sometimes used for Pedicel. ]'tiieil-sha]ied, the pencil (/xitii'i/liiin) being a brush or tuft of hairs. I'tniinti (-titiif}. Same as rinnate. !'< a a if" i- in (-urinif). In the form of a feather or its plume. /'( iiiiiin ITI il (-1 rrinf). Same as pin- nately nerved or veined; 93. /'<;//.(. ( ireek for live; gives compounds such as I', iiim-i'irjii llnri/. Composed of live car- l'< iiliii'lni iiiiiin. Name of a pentacarpel- lary fruit otherwise like a cremocarp. /'i ni, nli //limns (-us). With stamens in live clusters; -J-'iO. l'i ni,ir;i'/iiin. I.inn:eau artificial order characterized by l'< /'/"//// iioitg, i.e. five-styled flowers: :;:{~. GLOSSARY. 425 Pentdmerous (-us). Composed of five members in a circle; 176. Ptnta luli-iu. The Linnaean class with Pentandrous, i. e. five-stamened flow- ers; 249, 334. Pentapetalous (-us). Five-petalled ; 244. Pentaphyllous (-us). Five-leaved: 243. Pen/apterous (-us). Five-winged. Pentasepalous (-us). Of five sepals ; 244. /', n/ii.ftichous(-us). In five vertical ranks; 123. Pepo, Peponida, Peponium. A gourd- fruit; 298. Perennial (Perennis, Perennans). Last- ing year after year ; 32. Perfect (Perfectus). Said of a flower which is hermaphrodite. Pet-foliate (-at us). Where a stem seem- ingly passes through a leaf; 167. Perforate (-ittits). Pierced, or having translucent dots which look like holes. PergameneuS) Pergamentaceus. Parch- ment-like in texture. Peri. Greek for around; hence such compounds as Perianth (Perianthium). The floral en- velopes or leaves of the flower, consist- ing of calyx, corolla, or both ; 164, 243. Pericarp (-urpium). The fructified ovary; 286. Pericdrjric (-icus). Relating to the peri- carp. Perichcetial (-ialis). Relating to the Perickcetium, a set of bracts around the fruit-stalk in Mosses. Peridddium. The sheathing base of a leaf when it expands and surrounds the supporting branch. Perii-liiiiinii. Involucre of the capitu- ' lum of Composite ; 148. Per i< term (-ermu or -ermis). Outer bark or Epiphloeum. Periyone, Perigonium. Synonym of Perianth; 164. Perigynium. Name of hypogynous bristles, scales, or a sac, which sur- rounds the pistil (also the stamens when present) of many Cyperacese. Periyynous (-us). Literally around the ovary ; said of organs which are ad- nate to the perianth, or to this as con- nate with the low r part of the pistil; 182. Peripetalous (-us). Around the petals. Peripherie (-icus). Of or belonging to the circumference ; as of an embryo coiled round the outside of the albumen. Periphuranthiuiii. Synonym of the involucre of Composite; 148. Peripterus. Surrounded by a wing or thin border. Perisperm (-ermiurn). The albumen of the seed, at least the exterior and or- dinary albumen; 14, 310. Peristome (Peristoma or Peristomiitii/ ). The fringe or other structure surround ing the orifice (stoma) of a Moss. Peritropous (-us) or Peritropal. Said of a seed which is horizontal in the pericarp; or of a radicle pointed to- ward the sides of the pericarp. Persistent (-ens). Remaining even on the fruit, or over winter ; 243. Personate (-atus). Masked, as when a bilabiate corolla has a prominent pal- ate ; 248. Pervious (-ius). With an open passage- way. Perfuse (-usus). Having slits or holes. Perula, pi. Perulce. Scales of leaf-buds and the like; 40. Perulate (-atus). Furnished with pe.ru- lae or scales. Pes, gen. pedis. A foot. Hence in Latin compounds Longipes, long- stalked, Brevipes, short-stalked, &c. Petal (Petalum). A corolla-leaf; 165. Petaline (-inus), Petaloid (-oideus). Petal-like, or relating to petals ; 118. Pctalotly. Name for the metamorphosis of other organs (such as stamens) into petals; 174. Petiolar (-aris). Borne on or relating to a petiole. Petiolate (-atus), Petioled. Having a petiole. Petiole (Petiolus). The footstalk of a leaf; 85, 104. Petiolulate (-atus), Petiolular (-aris). Having a Petiolule (Petiolulus). A footstalk of a leaflet; 105. Pttraeus. Growing among rocks. Petrosus. Growing in stony places. Phcenogams, Phcenogamia, Pha'ii<>. A branch assuming the function of foliage : C.'i. /'/ii/lliiilirKdim (-< «.<). Delating to a Pliyllodium. A petiole usurping the form and function of a leaf-blade; 111). Phyllody, Phyllomorphy. Names for the transformation or metamorphosis of floral organs into leaves; 174. Phyllotdxis, Phyllotaxy. Leaf-arrange- iii. 'lit; 119. rin//li>iiiii>iiii. The unusual or abnormal production of leaves. piii/llojihiirr (-iiriuii ). The budding sum- mit of a stem on which leaves are de- veloping. J'lutl/iiin. Greek for leaf; 6, 85. See Phylla. Pkyllome, Phylloma. An assemblage of leaves, or of incipient leaves in a bud. Also recently used by German botanists for leaf generic-ally or poten- tially, that which answers to a leaf; 6. i/ritjihi/. Botany as relates to the de- scription and illustration of plants ; 345. PJiyldlogy. Synonym of Botany. Pliylumer, pi. Phi/lt'iim ni. Plant-ele- ments in morphology ; same as Plii/tdii. Greek name for plant; has been used in the sense of plant-ele- ment, or plant-unit ; 7. Plnjtolomy. Same as Vegetable Anatomy or Histology ; 2. I'ii-iuf. Pitch-black or brownish-black. I'ii-tii.t. Painted, or rather as if painted. Pileate(-ruii*. Having the form of a cap or Pilnts. I'i/i m-lii.-.ii. The mot-cap. J'ili/'i'i'iniit (-us). Hearing or tipped with linir.t (pili). Pilosciusculus. Slightly hairv. /'//n.-v (-n.fiia). Hairy, in general with any sort of pilo-ity; in particular with soft and distinct hairs. I'iiiiin. Our of the primary divisions of a pinnate leaf, either simply pinnate, when it is a leaflet, or a partial petiole or rhaeliis with the leaflets when the leaf is bipinnate; 104. Pinnate (-titus). When leaflets are arranged along each side of a com- mon petiole ; 100. ]'liiii (-it/in), PiftiUiftTiius. Said of a plant or a blossom provided with pistil, most properly for one having pistil only ; l!il. I'iflilliilinm. One of the names of the analogue of pistil in .Mosses, &c. /'i.-i/i/liH/i/. Name for the metamorphosis of other organs into carpels; 174. Pitcher. See Ascidium. A tubular or cup-shaped leaf, which usually holds some liquid ; 111. Pith. A central cellular part of a stem, especially of an exogenous stem ; 75. Pitted. Marked with small depressions or pits. Placenta, That in the ovarv which bears the ovules, sometimes the mere united margins of the carpel-leaves, sometimes a thickening or enlarge- ment of them, or even of some other part of" the ovary; 261. /'tiifi ittii/iun (-in). The disposition of the placenta-. Plin-i iitifurni (-«/•;» /.-•). Quoit-shaped, or in form like a flat cake. Plaited. See Plicate. Pliine (Planus). With flat surface or surfaces. Plutys. Greek for wide, in such com- pounds as Platyphyllus, broad-leaved, &C. Ph-'nix. Greek for full, used in com- pounds for several or many; as Tl< i<>- fihi'/lliiim, several-leaved, &c. Simi- larly /Yc/.f/as- for a great many. I'll ii i<-!i ii./i/t r, I 'ml if i rim). Bear- ing progeny, in the way nf otl'-lioots. J'nJi/'i rut it'll or f'rii/ijii-ii/iiiii is usually taken as the production by one organ of something different, such as the development of bmU and plantleN on leaver of le.ity shoots in place of (lowers, \-e. ; 73. /'rii/n/< run* (-//.<). Same as 1'roliferous. Prniii i I'ruiiii.--). Lying Hat. especially lace downwanl. Propdculum, l'i-"/ili i//<.<, J'ri'tii/ilii/tii. Alga, \-c.. the sii]iposed first jilants. Protoplasm, J'i-n/i>inliif. Wing-footed, i.e. petiole wing-margined, &c. rtifj-if. (ireek name for folding, as of leaves in a bud: I--'. L33. Pubi-iif. PttbeS I'sed for I'ube-cent. J'ulf ruin*. Minutely pubescent Pubnt. 1'tibes. eiice, hairinc". J'n/Hsctiit (-.».-'). Clothed or furnished with hairs or down, especially with soft or downv and short hairs. GLOSSARY. 429 Pufjioniform (-ormis). Dagger-shaped. Pullus. Dark-colored; dusky-brown or blackish. Pulvereus, Pulverulentus. Powdered ; as if dusted with powdery matter or minute grains. Pulvinate (-atus), Pulviniform (-ormis). Cushion-shaped. Pulvinus. A cushion ; name given to an enlargement or swelling close under the insertion of a leaf, or sometimes to the swollen base of a petiole. Puniilus. Low or little. Punctate (-atus). Dotted, either with depressions like punctures, or trans- lucent internal glands, or with colored dots. Puncticulate (-atus). Minutely punctate. Pungent (-ens). Terminating in a rigid and sharp point or acumination, like a prickle. Puniceous (-eus). Bright carmine-red. Purpureus. Originally the red of arte- rial blood ; but our purple is some- what dull red with a dash of blue or violet. Purpurdscens. Purplish. Pusillus. Very small, or weak and slen- der. Pustular, Pustulate (-atus), Pustulose (-osus). Having low elevations, like blisters. Putdmen. The shell of a nut; the endo- carp of a stone-fruit; 288. Pycnos, Greek for thick ; whence Pycno- cep/idlus, thick-headed, &c. Pyijmceus. Dwarf, pygmy. Pyramidal (-alls). Pj'ramid-shaped. Pyrene (Pyrena). Same as Nucule or Nutlet; one of the small stones of a drupaceous fruit : 2118. Pyretuirium, Pyridinm. A pear or pear- like fruit, same as Pomum. Pyrena i-ius. Name of a drupaceous pome, as of Medlar and Crataegus. Pyridion. Synonym of Pome. Pyrcnoan-p (-arpium). A general name for any drupaceous fruit; 292. Pyrif orni (-ormis). See Pear-shaped. Pyxidate (-atus). Furnished with a lid. Pyxidium, Pyxis. A capsule with trans- verse dehiscence, making a lid of the upper portion; 293. Quadri-. In Latin compounds, denotes four; as Quadrangular, Quadri/'ari- ous (in four vertical ranks), Quadriju- gate (in four pairs), £c. Quaternary, Quaternale. In fours or composed of four; 176. Quini, Quinary (-ius), Quinate (-atus). In fives ; 176. Quinque. Five. In Latin compounds, giving rise to such terms as Quincuncial, in a Quincunx; also five- ranked; 123, 136. Quinquefarious (-ius). In five vertical ranks. Quinquefoliate (-atus). Five-leaved. Quinquefoliolate, with live leaflets. Quintuple. Dividing into live parts, or five-fold. Quintujjlinerved or -veined. With mid- rib of leaf dividing into five (i. e. two lateral pairs) above the base; 93. Race. A variety of such fixity that it is reproduced by seed; also used in a looser and more extended sense for a series of related individuals without particular regard to rank ; 320. Raceme (Raccmus). An indeterminate or centripetal form of inflorescence with lengthened axis and equal-pedi- celled flowers; 146. Racemiferous. Bearing racemes. Racemifo'i-m (-ormis). In the form of a raceme. Racemose (-osus). Having the character or appearance of a raceme, or in ra- cemes. Raclds. See Rhachis. Radial. Belonging to the rav. Radiate (-atus). Spreading from or arranged around a common centre, or around the circumference of a cir- cle; bearing rays or ray-flowers. Radiately veined. Same as Palmately veined; 93. Radiatiform (-ormis). Said of a capitu- lum of flowers which is radiate by en- largement of some of the outer flowers, which however are not truly ligulate, as in species of Centaurea. Radical (-nlig). Belonging to or pro- ceeding from the root, or from a root- like portion of stem at or below the surface of the soil. Radicant (Radicans). Rooting. Radicel. A minute root or a rootlet. Radiciftorous (-its). Flowering (appar- ently) from the root. Radiciform (-ormis), Radicinus. Of the nature or appearance of a root. Radicle (-icula). Literally a diminutive root; but the "radicle " of the embryo, 430 GLOSSARY. so called in descriptive botany, is the hypocotyledonary and primal inter- node. (See < 'auliclc) ; 111. li'n/ii-ulose (-osus). Bearing rootlets. l['i the surface or epidermis, such as the chaff on the stalks of many Ferns. Ramification. Branching; 47. (-«*•). Flowering on the branches. -iw/jf). Branching or branchy. (-nuns). Bearing many branch- lets, i. e. Jtiiinuli or Ruphe. See Khaphe. lin/iliides or Rhaphides. Crystals in the cells of plants, especially needle- shaped crystals. Rdij (Radius). One of the radiating branches of an umbel (147); al.-o the marginal as opposed to the central part (or disk) of a head, umbel, or other flower-cluster, when there is a differ- ence of structure. Also used as an abbreviated expression for It'" i/-/l"/rers. Those which belong to the margin of a circular flower-clus- ter, and differ from (being usually large r than) those of the disk. Secuuli-scvncv. The adhesion of leaves or their stalks to a stem ; 158. Receptacle (JReceptdculum). A portion of axis forming a common support or bed on which a cluster of organs is borne. The receptacle of the flower, or the torus, is the axile portion of a blossom, that which bears sepals, petals, stamens, and pistils; 167, 211. The receptacle of inflorescence is tin- axis or rhachis of the head, spike, or other dense cluster; 143. l!< i-l'ilinlr (-i< rri/ifi. Straight-veined or straight- llerved ; !l'J. Hi ctin rial ( -in/if). Tn rectilinear ranks ; 124. li'i i-it i-ri'd (-ws), Recurvatus. Cunred backward or downward. Hi duplicate (-ntus) or RedupUcatimu. Folded and projecting outward. Reflexed (-us). Abruptly bent or turned downward or backward. Refracted (-us). Same as reflexed, but abruptly bent from the base. It'i i/ma. A two-several-lobed two- several-celled fruit (2-pluricoccous), which separates at maturity into as many -J-salved carpels, as in Euphor- bia : one form of hchi/.ocarp. A' i i/niiifiirji \-nrjiiiiiii). A general name of a dry and dehi.-ccnt Iruit, ^'J2. lit t.*). Inverted. Hi fiim tl!tt>is/i*). ^\'ith a shallow or obscure notch at a rounded apex : 97. Jtercrx/iiii. A changing back, or in the reverse direction ; 171. Renilntr (-ntiif). Rolled backward from the margins or apex; 133. Illiurlii*. The axis (backbone) of a spike or of a compound leaf; 101, 143. Rhnplte. The adnate cord or ridge which in an anatropous ovule con- nects the liilum with the chalaza; 279. 307. Rhipidium. A fan-shaped cyme; 156. GLOSSARY. 431 Rhizanthous (-us). Root-flowered ; flower- ing from the root or seeming root. Rhizina. The peculiar roots or root- hairs of Mosses, Lichenes, &c. Rhizocarpous (-us). Rhizocarpic (-icus). Literally root-fruited; used by De- Candolle for a perennial herb. Rhizome, Rlii:.u/nn. A rootstock; a stem of root-like appearance pros- trate on or underground, from which rootlets are sent off; the apex pro- gressively sending up herbaceous stems or flowering stalks and often leaves; 5G. Rhizomorphous (-us). Root-like in ap- pearance. Rhombic (-icus). Rhomb-shaped. Rhomboidal (-alls). Approaching a rhombic outline ; quadrangular, with the lateral angles obtuse. Rib. A primary and strong vein or conspicuous portion of the framework of a leaf ; 92. Ribbed. Furnished with prominent ribs. Rictus. The mouth or gorge of a bila- biate corolla. Rima. A chink or cleft. Rimose (-osus). With chinks or cracks, like those of old bark. Ring. In Ferns, &c. See Annulus. Rinyent (-ens). Grinning or gaping ; as is the mouth of an open bilabiate corolla; 248. Riparius. Growing along the banks of rivers, &c. Rivdiis. Growing along brooks. Rivularis. Growing in watercourses or rivulets. Root. The descending axis. Roots are axes which grow in the opposite di- rection from the stem, are not com- posed of nodes and internodes, are mostty developed underground, and absorb moisture, &c., from the soil; 27. Root-cap, 13, 28. Root-hairs. Attenuated unicellular outgrowths or hairs from the newly formed parts of a root, for absorp- tion; 13, 29. Rootlet. A very slender root or branch of a root. Rootstock. See Rhizoma: 56. Roridus. Dewy; covered with particles resembling drops of dew. Rosaceous (-eus). Arranged like the five petals of a normal rose; 246. Sometimes used for rose-color. Roseus. Rose-colored ; pale red. Rostellate (-atus). Diminutive of Ros- trate. RostMum. A diminutive beak. Also the name applied by Linnaeus to the Caulicle or Radicle. Rostrate (-atus). With a Rostrum, a beak or spur; narrowed into a slender tip or process. Rosular, Rosulate (-atus). Collected in a rosette. Rotate (-atus). Wheel-shaped; circular and horizontally spreading very flat ; 248. Rotund (Rotundus, Rotundatus). Round- ed in outline ; 95. Rouyh, Rouyhish. See Scabrous. RubMus, Rubescent (-ens), Rubens. Red- dish. Rubescent also is turning red. Ruber. Red in general. Rubicundus. Blushing, turning rosy- red. Rubiginose (osus). Brownish rusty-red. Ruderal (-alls). Growing in waste places or among rubbish. Rudiment. An imperfectly developed and functionally useless organ ; a Vestige. Rufous (-us), Rufescent (-ens). Pale red mixed with brown. Ruyose (-osus). Covered or thrown into wrinkles, Rugae. Ruminated (-atus). As if chewed ; said of the albumen of a nutmeg, &c. ; 311. Runcinate (-atus). Saw-toothed, or sharply incised, the teeth or incisions retrorse. Runner. A prostrate filiform branch which is disposed to root at the end or elsewhere ; 53. Running. Same as Repent. Rupestris, Rupicola. Growing on rocks or in rocky places. Rujitllis. Bursting irregularly. Rusty. Same as Rubiginose, Rufescent, and Ferruginous. Rutildiis. Deep red with a metallic lustre. Sabulosus. Growing in sandy places. Saccate (-atus). Sacciform. Sac-shaped; baggy. Sagittate (-atus). Sarjitlij'orm (-ormis). Arrow-head-shaped. Salsuginosus. Growing within reach of salt water. Salver-shaped. See Hypocraterimor- phous; 248. 432 GLOSSARY. Samara. An indehiscent winged fruit ; L'Ul. Samaroid. Resembling a samara. N ip-wood. New wood of an exogenous stem; 80. s,ii;-i,i;ir/> (-nr/iium). The succulent or Hc-liy piirtiiiu of a drupe; 285. Has been proposed a l-o as a general name for a baccate I'rnil : 292. Si/run iiti'ti (-usii.f). Producing long and lithe branches or runners, viz. Sunn, nt.< i Sarmi nin). Sativus. That which is sown or planted. S,nr-f, Hit/ml. See Serrate. Saxdtilis, Saxosus, Saxicolus. Living on or among rocks. Si-n/ii-ii/ii.t, Srii/irii/wulus. Roughish; diminutive of i/x S,-ii/t,T). Rough to the touch. ni (-ormis). Ladder-shaped; with transverse markings like the rounds of a ladder. Scales. Any thin scarious bodies, usu- ally degenerate leaves, sometimes of epidermal origin. Si- i//,,/n'it. Same as Crenate ; 98. Si-ii/i/. See Searious, Squamose. N, ••//// /,'//, /.--, 40. Si-inn/, Ht (-ens). Climbing, in what- ever mode; 51. Scape (Scapus). A peduncle rising from the ground; 51, 143. Scapiform (-ormis), Scapose (-osus). Re- sembling ;l scape. Scapigerous (-».-•). Scape-bearing. Scar. The mark left on the stem by the separation of a leaf, or on a seed, &c., by its detachment. Scarious or (-nrjiliini). \ pericarp which splits into one-seeded pieces; -_".tii. Si-ion. A young shoot : a twig used for grafting. Sriii ,-,,/,/, ,/.<. Like a si|uii-rel's tail. S,-/, i-'iiitl,'iiini. Name of the fruit of -Mirabilis, and the like; an akene (']" losed in an indurated portion of calyx-tube. Scleroideus. Having a bard texture: from Si-/, /;,.<, hard. Si-,,/,;/',,,-,,, (-ormis). Having the appear- ance of sawdii-t. Scorpioid. \ form of unilateral inflo- rescence whieh is circinately coiled in tlie bud: in the stricter sense, a f.inn with tlie ilo«er< two-ranked, these beiiiLT thrown aliernatelv to the right and left : l.-)5. 157. Scrobiculate ( atii?). Marked by minute or shallow depressions. acrotifortn (-"/•/;//.<). Pouch-shaped. Scurf. Small and bran-like scales on the epidermis. S, -at, it, (-ntu.<). Si-H/ifurm (-ortnis). Buckler-shaped. s,-,itilHj'<,,-,ii (-,,/•///!.<). Platter-shaped. Sfymetnr-gliiijitd. See Acinaciform. Sectile (-ilia). As if cut up into portions. S,ition (Sectiu). In classitieation, is applied in a general way to a divi- sion in the arrangement of genera. Species, or .,ther Croups ; 327. Sectus. Completely divided; 99. Secund (Secninlux). When parts or organs are all directed to one side. Secundiflorus. With flowers of a cluster all secund. Secundine. The second (inner) coat of an ovule; 277. Seed. The fertilized and matured ovule ; the result of sexual reproduction in a phamogamous plant; 305. Seed-leaves. Cotyledons, 11. Seed-stalk. See Funiculus and Podo- sperm. Ni i i/-n we/. See Pericarp. Setjftdlis. Growing in grain-fields. Ni ,/inent (Snjn,, niinn). One of the divisions into which a plane organ, such as a leaf, may be cleft. Segregate (-atus). Separated; kept apart. N, n/i n. Seed. N« /,//. Hah', in Latin compounds : such as Semi-adherent. The lower half adhe- rent, (S;c.; Ni ////-, n, if, li.rirniil (-inili.-i), half clasping the stem ; Semiovtit*, ovate halved lengthwise. &»•. Semiandtropous. Same as Amphitro- N, inilintiir, Si'iniliiiiiiti (-ntn.<). A svn- onym of Lunate, being like a half- moon. (-nllf). Relating to the seed. N, niinij i runs (-».. Ni /ml ( Si'/inlm/i). A calyx-leaf: li'i.'i. S, fulfill,' (-inn."). Si/minim. Relating to sepals. Ni /ml, ,1,1 (-,,1,1, US). Kesemblinij a sepal. Ni /I.I/IH/I/. Name for the metaniorpho-js of petals. &C.. into sepals or sepaloid organs: 174. S, /nil-lit, i/ itii/ri /•.<. Those of distinct sexes; same as Diclinous; 191. GLOSSARY. 438 Septate (-atus). Separated by a parti- tion or septum. Septicide, Septicldal (-cidus). When a capsule dehisces through the dissepi- ments or lines of junction ; 289. Septi/'erous (-us). Bearing the partition or dissepiment. Septifrayal (-us). Where the valves in dehiscence break away from the dis- sepiments ; 290. Septum. Any kind of partition, whether a proper dissepiment or not. Septulute (-titim). Divided by spurious or transverse septa. Serial (Serialis) or Seriate (Seriatus). Disposed in series or rows, whether transverse or longitudinal. Sericeous (-eus). Silky; clothed with close-pressed soft and straight pubes- cence. Serotinous (-us). Produced compara- tively late in the season. Semite (-atus). Beset with antrorse teeth; 97. Serrulate (-atus). Serrate with very small or fine teeth; 97. Sesqui. A Latin prefix denoting one and a half; as, Sesquipedalis, a foot and a half. Sessile (-His). Sitting close, without a stalk ; destitute of peduncle, pedicel, or petiole, as the case may be. Seta. A bristle, or bristle-shaped body. Setaceous (-eus). Bristle-like. Setiform (-ormis). In the form of a bristle. Setiyeroiis (-us). Bristle-bearing. Setose (-osus). Beset with or abounding in bristles ; bristly. Setula. Diminutive of Seta. Setulose (-osus). Bearing or consisting of minute bristles. Sex. Latin for six; as in Sexangular, Sexfarious, Sexpartite, &c. Shayyy. Pubescent with long and soft hairs ; same as Yillous. SJieath. A tubular or enrolled part or organ, such as the lower portion of the leaf in Grasses. See Vagina. Sheathing. Enclosing as by a sheath. Shield-shaped. In the form of a buckler; plane and round or oval, with stalk attached to some part of the under surface ; 96. See Clypeate, Scutate, Peltate. Shrub. A woody perennial of less size than a tree : 50. Shrubby. Havingthe character of a shrub. Sieve-cells. 77. Siyillate (-atus). As if marked with the impression of a seal, as the rootstock of Polygonatum. Sigmoid (-oideus). Doubly curved like the Greek s or the capital S. Silicle (Silicula). A short silique, not very much longer than wide ; 294. Siliculosa. Name of the Linmean arti- ficial order of the class Tetradynamia, having Siliculose pods ; 337. Silique (Siliqua). The peculiar pod of Cruciferse, especially when much longer than wide ; 293. Siliquosa. Name of the other order of Tetradynamia, with Siliquose fruit, i. e. a Silique ; 337. Silky. See Sericeous. Silver-grain. The glittering plates in exogenous wood belonging to the medullary rays; 74. Simple (Simplex). Of one piece, series, &c. A simple pistil is of one carpel ; a simple leaf, of one blade, &c. Simple Fruits, 291. Siinplicissimus. Most simple; complete- ly simple. Sinatrdrse. Turned or directed to the left ; 51, 140. Sinuate (-a,tus). With a strongly wavy or recessed margin ; 98. Sinus. A recess or re-entering angle. Slashed. Same as Laciniate. Smooth. Either opposed to scabrous, i. e. not rough, or to glabrous, i. e. not pubescent ; the former is the more correct application. Soboles. Shoots, especially those from the ground. Soboliferous (-us). Bearing vigorous lithe ?hoots. Solid Bulb. A corm ; 61. Solitary (-arius). Single, only one from the same place. Solubilis. Separating into portions or pieces. Solutus. Loosed ; becoming separate. Sordidus. Of a dull or dirty hue. Sorediate (-atus). Bearing small patches on the surface. Sorema. A heap of carpels belonging to one flower; 263. Son', sing, sorus. Heaps, such as the clustered fruit-dots of Ferns. Sorose. Heaped or bearing Son. Sorosis. A fleshy multiple fruit, such as a mulberry, bread-fruit, and pine- apple. Spadiceus. A bright and clear brown, or chestnut color 434 GLOSSARY. f. Having the nature of or bearing a ii.r. A spike with a fleshy axis; 149. . The length of the space between the tip nl tin- thumb and that of the little linger, when outstretched : about nine inches. s. Sparse or scattered; whence .»-, with scattered flowers; Sparsifolii/*, with scattered leaves, &c. Sjiiit/nn-tniix (-us). Spathe-bearing, or of the nature of a S/HI//I, (Spilt/in). A large bract, or a pair of bracts, enclosing a flower-clus- ter; 142. Spatliellu. An unused name for the glumes of Grasses. Spathilla. A secondary or diminutive Bpathe. Spiilitlnte (Spathulatus). Oblong with the lnwer end attenuated, shaped like a druggist's spatula; 95. Species. The particular kind, the unit in natural history classification; 317. Specijic (,'lniriii-li r. A"/'//.•. i>'/<>n,/it>l< (-lulu). Name given to young root-tips; once sup- posed to be a peculiar organ; 28. Sporadic (-icus). Widely dispersed or scattered. Sporangium. A spore-case or theca con- taining the analogues of seeds (spores) in the higher Cryptogams. Spore (Spora, Greek for seed). The analogue of seed in Cryptogams. Spore-case. See Sporangium. ^piiridiuin. Synonym or diminutive of Spore. .sy;«,v/'< naif. Spore-bearing. ^jii'ifiK-ni-p (-in-plum). Name given to certain spore-cases, as of Lycopodi- aceae. fyiiirvphore (-ontm). One of the syno- nyms of Placenta. Xjmriilt (S/n'n-iiln). Diminutive spore or a sort of spore. >/j. //•«//'/'. runt (-us). Bearing or con- taining spores. >'/ii'/-f. A bud-variation or seed-varia- tion; 319. Xpmiii.ai-1 ni (-1 ii.<). S/iiinnifc. Froth-like in appearance. Spur. A hollow and slender extension of some portion of the hlo-som. 11-11- ally nectariferous, as of the calyx of Larkspur and the corolla of Violet: rarely applied also to a solid spur-like 'process. S/inrr,'/. Producing a spur. See Cal- carate. Squama. A scale of any sort, usually the homologue of a le.it. Squnmate (-ntu.t). Squamiferous, Squa- musiis. Furnished with scales. GLOSSARY. 435 Si/iiamella, Squdmula. Diminutive squama; scales of secondary order or reduced size. Si/iiii/iii/'uriii (-iii-mta). Scale-like. Squnmulose (-osus). Covered or beset with minute scales. Squarrose (-osus). Literally rough- scurfy; applied to bodies rough with spreading and projecting processes, such as tips of bracts, &c. Squarrulose (-osus). Diminutively squar- rose. Stni-lii/K. Greek for spike. Stalk. Any kind of lengthened support on which an organ is elevated. Stamen. One of the elements or phylla of the andrcecium ; 105. Stamineiil, Stamineaus (-&us). Relating to the stamens ; 191. Staminiferous (-us). Stamen-bearing. StaminotHum. A sterile stamen, or what answers to a stamen, whatever its form, without anther. Staminody. Name for the metamor- phosis of other floral organs into stamens ; 174. Standard. The posterior petal of a papilionaceous corolla; 184. Stuns. Supporting itself in an erect position. Station. Particular place as to soil, ex- posure, &c. , which a plant affects ; 366. Stellate (-atus). Star-shaped, arranged like the rays or points of a star. Stellulate (-atus) or Stellular. Dimin- utive of Stellate. Stem. The main ascending axis of a plant; 45. Stemless. See Acaulescent ; with no leaf- bearing stem above ground; 45. Stemlet. Diminutive stem ; as that of the plumule. S/CHOA-. Greek for narrow ; hence N/r inijiliijllus. Narrow-leaved, &c. S/i rii/nia. Any foliaceous prolongation of the blade of a leaf down on the stem by decurrence. Steriymitm. Name of Desvaux for the Dieresilis of Mirbel. Sterile (-///.-.•). Barren, as a blossom destitute nt pistil, 191; a stamen without anther, or an anther without pollen; an ovary, without good ovules, seeds without embryo, &c. In com- mon English use, a male or staminate flower is said to be a sterile flower. Stichus. Greek for row or rank, usually meaning vertical rank ; hence such compounds as Distichous, two-ranked ; J'/i.-ii.<. three-ranked, &c. Stiyma, pi. stiymata. That part or sur- face of a pistil (usually on or a part of the style, or in place of it) which re- ceives the pollen for the fecundation of the ovules ; 166. Stly inn tic (-icus), Stiymntose (-osus). Relating to stigma. Stigmatiferous. Stigma-bearing. Stings. Stinging hairs, seated on a gland which secretes an acrid liquid, as in Nettles. Stipe (Stipes). A stalk of various sorts; the support of the cap of a mush- room ; the leafstalk of a Fern ; any stalk-like support of a gyncccium or a carpel; 212. Stipel (Stipellum). An appendage to a leaflet analogous to the stipule of a leaf; 106. Stipulate (-atus). Provided with stipels; L06. Stipitate (-atus). Having a stipe or special stalk. StipitiJ'nrm (-ormis). Shaped like a stipe; stalk-like. Stipulaceous (-eus), Stipular (-aris). Belonging to stipules. Stipulate. Possessing stipules. Stipules. Appendages or adjuncts of a leaf one on each side of the insertion ; 85, 105. Stirps, pi. stirpes. A race. Stock. Synonym of Race ; also the portion of a stem to which a graft is applied; a caudex, rhizoma, or root- like base of a stem from which roots proceed; 51. Stole, Stolon (Stolo). A sucker, runner, or any basal branch which is disposed to root ; 53. Stolontferous (-us). Sending of or propa- gating by stolons, runners, &e. Stomn, pi. stdmata, Stomate. One of the apertures in the epidermis of folia- ceous parts, through which cavities within communicate with the external air; 89. Stomatiferous (-us). Bearing stomata or "breathing pores." Stone. The hard endocarp of a drupe. Stone-fruit. A Drupe, such as a peach or plum; 297. Stool. The plant from which layers are propagated, by bending down to the ground to be rooted. Stramineous (-eus). Straw-like or straw- colored. 436 GLOSSARY. Strap-shaped. See Ligulate (247) and l.i i rate. Strinii (-ut us). Marked with fine longi- tudinal lines, streaks, or diminutive grooves or ridges (Stria). Strict (Strictus). Close or narrow and upright ; very straight. BtriffUlose (-osus). Minutely strigose. Strii/usi (-n.nif). lieset with strii/ie, or sharp-pointed and appressed straight and stiff hairs or bristles. St robilaceous (-i-us), Strobiliform (-or- in'tK). Relating to or resembling a Strobile. Strobile (Strobilus). An inflorescence formed largely of imbricated scales, as that of Hop and a Fir-cone; 303. Stro'nbuliformis, Strombuliferus. Twist- ed spirally into a screw shape, as the legumes of the Screw-bean (Proso- pis, sect. Strombocarpa) and of some species of Medicago. Stropliiole (-tola). An appendage at the hilum of certain seeds; 308. Structural Botany, 2. Struma. A wen or any cushion-like swelling on an organ. Strumose (-osus), Strumiferous (-us). Furnished with a struuia or goitre-like swelling. Stupose (-oats). Tow-like: with tufts or mats of long hairs. Stylt ( Sty/u*). The usually attenuated portion of a pistil or carpel between the ovary and stiguia; 16G. Styliform (-ormis). Style-shaped. Stijliferous. Style-bearing. Stylinus. Belonging to the style. With styles of remarkable or number, &c. Stylopodium. An enlargement or a disk- like expansion at the base of a style, as in rmlicllil'ene. K/I/I. In c-dinposition of Latin words in terminology, denotes somewhat or slightly ; as. Sii/im-nti , S/i/iruniutt , that i> aeutish. somewhat cordate, &c. in-iilni, and Siiliini/trirate in aestiva- tion, i::r. >'///« r«.«•). As if cut or broken off at the lower end. Succubous (-us). When in leaves crowded on a stem the apex of eac*i leaf is covered by the base of the next above. Succulent (Xuccosus). Juicy. Sucktr. A shoot of subterranean ori- gin; 53. Su/ruttsctnl (-ens). Slightly or ob- seurely shrubby; 50. Kujl'i-iitu: An undershrub. Suffruticose (-osus). Low and shrubby at base; 50. Sujf'ultus. Underpropped or supported. Sulctitr (-fittis). Grooved or furrowed. Super. Above. See Supra. Superior, Superus. Growing or placed above; also in a lateral flower on the side next the axis ; thus the poste- rior or upper lip of a corolla is the superior; lljO, 183. Superposed (Superpositus). Verticallv over some other part. Knpt rjH mi/ luii, 179, 195. Supervolute (-?/.«), Kiipc-iivlutire (-ivus), Same as Convolute when applied to plaits; 139. Supine (-inn.-!). Lying flat with face up- ward. Xi'/>j>ression. Complete abortion; 179, 190. Xti/ira. Above ; hence in Latin com- pounds, Supra-axillary, above the axil; 8«prq/b/fac€ptis,above a leaf, &c. Supradedecompound. Several times com- pound. Surculos< (-IISHS). I'rodncin.ir suekers. Surculvs. A sucker; a shoot rising from a subterranean base; 53. Kursinii. I'pward; directed upward or forward. Suspended (N//s/-, ».«».<). Hanging di- rectly downward: hanging from the apex of a cell. •s'»7" "-•'"' of the embryo, 284. Siitin-,1/ (-ii/is). ];,. latin- to a suture. Suture (-ura). A junction or seam of union : used commonly as a line of opening; 260. Sword-sJirrped. A blade with two sharp and nearly parallel edges, as in Iris. GLOSSARY. 437 Syconium or Si/conus. A multiple fruit like that of tin- Fig; 148, 303. ,^1/lri.slris. Growing in woods. Symmetrical. Regular as to number of parts or as to shape. In the blossom it denotes the former; 175. Symmetry. In the flower relates to symmetrical disposition of organs on the axis ; 174. Sympetalous (-us). With united petals; same as Gamopetalous ; 244. Symphiantherous (-us). Same, as Sy- nantherous and Syngenesious. Symphysis. Same as Coalescence. Symphystemonous. \\'ith stamens united. Sympode, Sympodiitm. A stem made up of a series of superposed branches in away to imitate a simple axis; a Sympodial stem ; 55, 154. Synacmy. Same as Synanthesis. Synantherous (-us). Stamens coalescent by their anthers. Synanthesis. The simultaneous anthe- sis or readiness of the anthers and stigmas of a blossom ; 219. Syncarp, Syncarpium. A multiple fruit such as a mulberry, or a fleshy aggre- gate fruit, like that of Magnolia; 299. Syncarpous (-us). Composed of two or more united carpels; 261, 263. Syncotyledonous. With cotyledons sold- ered together. Synedral. Growing on the angles. Synema. The column of monadelphous filaments, as in Mallow. Synaenesia. Linna?an class (335) charac- terized by having the anthers united or Syngenesious. With anthers cohering in a ring; 250. Synonym. A superseded or unused name ; 354, 365. Synonymy. All that relates to syno- nyms ; 365. Synsepalous (-us). Of coalescent sepals ; same as Gamosepalous ; 244. Systematic Botany, 2. Systylus. The coalescence of styles into one body. Tabescent (-ens). Wasting or shrivel- ling. Tail. Any long and slender terminal prolongation. Taper-pointed. See Acuminate. Tap-root. A primary descending root forming a direct continuation from the radicle : 31. Tawny. Same as Fulvous-, dull brown- ish-vellow. Taxolor/y, Taxonomy. Relating to clas- silieaticm and its rules; 3, 315. Teeth. Any small marginal lobes. Tvymtn. The inner coat of a seed; 306. Telu. Latin name for tissue, cellular tissue, &c. Teleianthm. Same as perfect, or her- maphrodite-dowered. Tendril. A filiform production (either axile or foliar) by which a plant may climb ; 54. Tepal ( Tepalum). A division of peri- anth, whether sepal or petal (hardly ever used). Teratological. Relating to malforma- tion or monstrous conditions. Teratology. The science of monsters and malformations ; 170. Terete (Teres). Round in the sense of having a circular transverse section. Tergeminate (-atus). Thrice twin. Terminal (-alis). Proceeding from or belonging to the end or apex ; 7. Terminology. Same as Glossology ; 3, 359. Ternary (-arius). Same as Trimerous ; consisting of three; 176. Ternate (Ternus, Ternatus). In threes; as three in a whorl or cluster. Tessellated (-atus). In chequer-work. Testa. The outer seed-coat, which is commonly hard and brittle, whence the name, which answers to seed-shell ; 305. Testaceous (-eus). Of the color of un- glazed common (brownish-yellow) pottery. Tetra. In Greek compounds, four ; hence Tetracdrpellary (-aris). Of four iar- pels; 261. Tetracdmarous (-us), Tetracoccus. Of four closed carpels. Tetradyndmia. Linnsan class (;;.'!,">) which has the stamens. Tetradynamous (-us). With four long and two shorter stamens ; 250. Tetragonal or Tetragonous (-us). Four- angled. Tetraf/ynia. Linnaean artificial order (337), characterized by having the gynoecium. T( trdgynous. Of four carpels or styles. Tetrdmerous (-us). Composed of four members in a circle; 176. Tetrandria. Linn;\?an class having the flowers perfect and Teirnndrous. With four stamens; 249, 334. GLOSSARY. Tetmpetalous (-us). With four petals; 244. / , !r,iolii/lloUS (-US). Four-lij:i\ ril ; ^ 1 '•',. T< ti-oijin Iron* (-11.1), With four sharp in- -alinit angles. Titrn.ii /in/, ,11.1 (-u.i). \\'ith four sepals; 244. Tttni.itirliiius (-us). In four vertical ranks. Tkiilniiiijloroiif (-iff). '-',W. \\'ith parts of tin- llower hypogynons, nr on the T/iiil.iiiiii.i. The receptacle of a flower: Ki7. See Toms. Tliiilli>j>lii/ti.i \ Thallophyta), :341. Timlin*. \ -i nit urn, in place of Mem anil folia;,' , Tlu-ca. A case; an anther-cell (251); a spore-ca.-e, \c. (An early name for the anther, 1GG.) TliK-iiji/ii.'i'i (-orum). The stipe of a carpel (homologous with petiole); 212. Thorn.. Same a- spine; 55. Tlnil.i>.ie (-onus). Diminutively or slightly torose. Tortus. Twi-ti-d. Tortilix. Siisceptil.Ie of twisting. Torus. The receptacle of a flower; 167, 211. Trabeculate (-atus). ('m-— liarred. Trachea. A spiral vessel or duet, named from resemblance to the tracheae of insects. 'J'/-io-/ii/i-iir/ion.i (-us) Hough-fruited. Trachyspermous. Rough-seeded, \-p. T i-on.ii-i r.if (-trsus). Across; right and 1. It a> to bract and axis; collateral; 160. Triij»r.ij'orm (-ormis), Trapezoid. Cn- symmetrically four-sided, like a tra- pezium. '/'/•( t. A woody plant with an elevated trunk. Tri-. In compound words, both Latin and Greek, denotes three or triple. Triachcmium. A fruit like a eremocarp but of three carpel>. Trtmkljiltoiis, Trinlti-/>o>i.i (-u.i). Consisting of three fruits or carpels. f (-H.I). Bearing three heads. oiix (-11.1). Hairy -fruited. Tr'n-lioilt.i. IJesemhling hair. Trii-l,,'iiomous (-us). Three-forked; branched into three divisions. Ti/i-lionn: (Tricliomn). Any outgrowth of tlie epidermis, such as a hair or bristle; 209. TricM-ruus (-us). Consisting of three cocci. Tricolor. Three-colored. Trirusjiiiliiti- (-a /us). Tipped with three cusps or pointed tips. (-ntu.i). Three-toothed. i'. Thrice digitate. Triiluu.i. Lasting for three days. Ti-'n in/in/ ( Trif-nnis). Lasting for three years. Tr'iforioiiK (-hif). Facing three ways, in three vertical ranks. Tr'iji«////. See Lanate and Tomentose: clothed with long and tortuous or matted hairs. Xanthos. Greek for yellow in com- pounds, MU-II a> .\/>!j'j>- That which can be bisected in only one plane into similar halves; IT-'i. ADDENDA. Antidronwus, Antidromy. When the course of a spiral is reversed, 157. Infertile {-His}. Said of a pistil or flower which fails to set fruit. I'nli/tmbryony. The production of two or more embryos in a seed, 284. Saprophytes (-yta). Plants feeding upon decaying vegetable or animal matter