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GRAY’S BOTANICAL TEXT-BOOK. 


VoLumME I. 


STRUCTURAL BOTANY. 


GRAY’S BOTANICAL TEXT-BOOK 


CONSISTS OF 


Vou. I. Srrucrurat Borany. By Asa Gray. 
II. PuysioLoeicat Botany. By Grorce L. Goopate. 


Ill. Intropuction To Cryprocamic Botany, BOTH 
o 
StrucTURAL AND Systematic. By Wi iiam G. 


Fariow. (Jn preparation.) 

IY. Sxetcu or THE NatTuraL OrpDEers 0F PH£NOGAMOUS 
Piants; their Special Morphology, Useful Pro- 
ducts, &c. (Jn preparation.) 


au 


GRAY’S BOTANICAL TEXT-BOOK. 


(SIXTH EDITION.) 


Wor. IE. 


SrRUCTURAL . BOTANY 


OR 


ORGANOGRAPHY ON THE BASIS OF 


MOA? TOLOG YY: 


TO WHICH IS ADDED THE PRINCIPLES OF 


TAXONOMY AND PHYTOGRAPHY, 
AND 


A Glossary of Botanical Terms. 


BY 
ASA “GRAY, LL.D., Ere., 


FISHER PROFESSOR OF NATURAL HISTORY (BOTANY) IX 
HARVARD UNIVERSITY. 


IVISON, BLAKEMAN & COMPANY, 


PUBLISHERS, 
NEW YORK AND CHICAGO. 


Copyright, 
By Asa GRAF, 
1879. 


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 Phznogamous 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 Phznogamous 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. 


1v PREFACE. 


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 fourth volume, a sketch of the Natural Orders of 
Phenogamous 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, 
CamBripGe 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 
Y INTRODUCTION. THe DEPARTMENTS OF THE SCIENCE 1 


CHAPTER I. OUTLINES OF THE GENERAL MORPHOLOGY 
OSE HE NOGAMOUS ELANTS) = 0s ef 2 21s 6 5 


CHAPTER Il. MORPHOLOGY AND DEVELOPMENT OF THE 


EAB RVOSAND. SEE DEUNG: 2.0 ee cn) een © 9 
The Embryo, its Nature, Structure,and Parts . . ... . 9 
Development of the Dicotyledonous Embryo in Maple . . . 10 
In Ipomea, or Morning Glory, &c., with Albuminous Seeds . 13 
In Embryos with thickened Cotyledons . . . ..... 16 
As of Almond, Beech, Bean, &e.. . . . 17 


With Hypogeous Germination and no Elongation ‘of Grable 19 
In Megarrhiza, &c., with concreted Petioles to the Cotyledons 21 
In Ipomea leptophylla with foliaceous and long-petioled 


Cotyledons and no elongation of Caulicle. . . . . . . 22 

In Pumpkin, &c., with no Primary Root . ....... 2 
The Polycotyledonous PMDLY Ones ee 23 
The Monocotyledonous Embryo of Iris, Onion; ‘Cereal Gane 24 
_ Pseudo-monocotyledonous and Acotyledonous Embryo. . . 26 
Dicotyledonous and Monocotyledonous Plants . . . . . - 27 


CHAPTER II. MORPHOLOGY AND STRUCTURE OF THE 


ORGANS OF THE PLANT IN VEGETATION .. 27 
PSECHLON La ORC THE ROOTS ToL: pr lise he spades fuss le Va ea oll PL 
Nature, Growth, and Compstition mera tee eh EDS 
LSYGTA TERNS Berean Oe ati aes 8 Me ite A ie ale lah ee Mey RI 
BITIASFOMMOOLSteeMC nin ons I ncas ee cal ee eee Atos See eee 
Dna OncmATTIaIse a cs Eacsa- tu eo eas os to ek ee. 80 
ICH DIGae Res ee Subse s Sol APC. FN ts mares OR 
IBCKenni ashy wbowe See ty ub) ceca ee oe at eee a Be 
Aerial Roots shies Ba ey oie ey et A a eT a es, Ee 
Epiphytes or Air- Seilasile aswel. 5 Se Thee ape ks 


Parasitic Plants, Green and Colored He Aap ee ee Fe orn Os 


CONTENTS. 


Section II. Or Buns 


Scaly Buds and Bud-seales . : 
Naked, Subpetiolar, and Fleshy Buds : 
Bud-propagation : 
Norinal, Accessory, and desi: Buds : 


Section III. Or tHe Stem 


§ 1. GeNERAL CHARACTERISTICS AND GROWTH 


Development and Structure 
Ramification, Branches ‘ 

Excurrent and Deliquescent Stems : 

Definite and Indefinite Annual Growth 


§ 2. Forms or STEM AND BRANCHES 


Herbs, Shrubs, Trees, Culm, Caudex, Scape 
Climbing Stems, Twining or otherwise 
Leaf-climbers, Tendril-climbers, and Rept-elarbers 
Suckers, Stolons, Offsets, Runners 

Tendrils formed of Stems F 
Sympodial and Monopodial Stems 

Spines or Thorns and Subterranean Stems 
Rhizoma or Rootstock 

Tuber, Tubercles 

Corm or Solid Bulb 

Bulb, Bulblets 

Condensed Aerial Semen 


Stems serving for i Phyllocladia, » Cladophy lla 


Frondose Stems . 


§ 3. INTERNAL STRUCTURE 


Anatomical Elements . 

Endogenous Structure 

Exogenous Structure ; its Besiaiane 

First Year’s Growth 

Pith, Layer of Wood, &e. 

Bark, its Parts and Structure 

Annual Increase in Diameter . 

Demarcation of Annual Layers 

Sap-wood and Heart-wood . 

Growth and Duration of Bark * : 
Living Parts of a Tree or Shrub, Longevity . 
The Plant composite . 


Section IV. Or LEAVES 


§ 1. Tuerr NaTuRE AND OFFICE 


Parts of a Ileaf . 
Duration, Defoliation, Normal Beacon 


CONTENTS. 
§ 2. TuHerr SrrucTuRE AND Forms As FOLIAGE . 


Internal Structure or Anatomy 

Parenchyma-cells 

Epidermis, Stomata or roving: pores 

Framework, Venation . . Sa ees 

Parallel-veined or Nerved ee CS er ween Bat So) ne 

Reticulated or Netted-veined Leaves 

Pinnately or Feather-veined and Palerdtely or adiately 
Veined aN ED 

Honma as tonOutdiney oye vo es ke (oss es ee Ee 

Forms as to Extremity ; 

Forms as to Margin or Special G@eadine and Dentekon - 

ihobationvor/Seomentation «2 4 2s. | 2) 20's. ae 

Number and Arrangement of Parts . 

Compound Leaves, Pinnate and Palmate or Dipitate &e: 

Petiole or Leafstalk 

Stipules, Ligule, Stipels . 

Leaves in unusual Modifications . 

Such as Inequilateral, Connate, Periolate 

Vertical and Equitant 

Without distinction of Parts 

Stipules serving for Blade 

Phyllodia, or Petioles serving for Blade 


§ 3. LEAVES SERVING SPECIAL OFFICES 


Utilizing Animal Matter 
Ascidia or Pitchers 
Sensitive Fly-traps 

Leaves for Storage : 
Bulb-scales and Bud-scales . 


CHAPTER IV. PHYLLOTAXY, OR LEAF-ARRANGEMENT 


Section I. DistrisuTion or LEAVES ON THE STEM 


Phyllotaxy cither Verticillate or Alternate, Cyclical or Spiral 
Verticillate or Cyclical Arrangement 

Alternate or Spiral Arrangement 

Its Modes and Laws : 

Relation of Whorls to Spirals . 

Hypothesis of the Origin of Both 


Fascicled Leaves 

Section IJ. Disposition or LEAVES IN THE Bup . 
Vernation and Mstivation; the Modes .......4. 
Direction. Dextrorse and Sinistrorse ....... 


Vili CONTENTS. 
CHAPTER V. ANTHOTAXY OR INFLORESCENCE... . 141 


Bracts and Bractlets and their Modifications . . .. . . I4l 


Peduncles, Pedicels, Rhachis, Receptacle ..... . . 143 
Position of Flower-buds, Kinds of Inflorescence ... . . 144 
Indeterminate, Indefinite, or Botryose. . . ... . . . 146 
Raceme, Corymb; Umbel) 2 5) 3) ae) Sec a 
Meador Capitultnige-aysee nae Reet Go 1Gt/ 
Syconium or Hypanthodium ..... . . 9. «. » «= «= 148 
Spike, Spadix, Ament or Catkm .©. .-. «= % 205 =e 
Panicle and other Compound Forms ....... . . 180 
Determinate or Cymose.= -.-  @ a) =a 
Cyme, Glomerule, &. . . Mera an sc LE 
Botryoidal Forms of Cymose Ty oe | sts sta: Sap shea ee ee 
Sympodial Forms. . . 154 
Scorpioid and Helicoid, the Pinatas oe “4 
Monochasium . . PR ESS 
Bostryx, Cincinnus, Epadine i omen ‘&e Me re EY 
Mixed Inflorescence =<. «i+ 2: By = & 4. tes 6) eens 
Thyrsus, Verticillaster,&c. . . a: rb nales) PET 
Relations of Bract, Bractlet, and lower “oe eu pt ee 
Anterior and Posterior, or Inferior and Superior . . . . . 160 
Mediamand: ‘Transverse! = <=) q: «) o<l 6) oscuro 
Position:of Bractlets: sn. Gy a a) a te oe ee 
Tabular View of Inflorescence .. - . . 2 « %  » « =» aioe 
CHAPTER VL THE FLOWER . =. ..>< =. =. ee 
Section I. Irs Nature, Parts, anpD MeTaAMoRPHY .. . . 163 
Floral Envelopes, Perianth, or Perigone . . ..... . 164 
‘The Parts, Calyxsandi@orolla, 7 <2) sew 2) eee 
Androecium, Stamensy.° . <=. = = -gen. (. 1) aeons 
Gynecium; Pistils.-y 3. oe iy a ie Le ee fe eee 
Torus or Receptacle of the Flgsres oy! eo joc ta ee 
Metamorphosis . . . oe) 3o- OI 
Unity of Type cient i Posae oad Fcsuasititners «ee ED 
Teratological Transitions and Changes . ...... . 170 
Srorion, Dk, Bio ran, SyMweRRY 2. % 2.0 2) <5 2 ele ee ee 
Symmetrical, Regular, and Complete Flower . . .. . . 1785 
Numerical Ground-plan-.) <  : = < =o. ssi, 2) sures 
Pattern. Flowers’02 1. 8 0s gen so sae) ee 
Diplostemonous Type... <= =... sy ea 
Section III]. Various MopiricaTions OF THE FLOWER. . . 179 


§1. "ENUMERATION Or THE KINDS -. 2% =<) < =i. seseeeeeee 


§ 2. Recurar Union or Srmiuar. Parts ..-. ... . 180 


Coalescence or'Cohesionicus 60 )sn0) ame 2. eee) eee eee 


CONTENTS. jie. 


§ 3. Union oF DisstMILar OR SuccESSIVE Parts . . . . 181 
Adnation or Connation . . Cer Re ren oan vated oh LOD 
Hypogynous, Perigynous, papers NG Te RCE aah lee iceaar ia tS; 

§ 4 I®REGULARITY OF StmmtaR Parts ....... . 184 

§ 5. DrisAPPEARANCE OR OBLITERATION OF Parts . . . . 187 
Abortion or Suppression of Parts of a Circle . . . . . . 187 
Abortion or Suppression of whole Circles. . . . . . . . 190 
‘Lemus therewith:conneeted) | Eos .0 so ee eee «(LOL 
Suppressed Perianth . .. . Te yc oe eee op BLO 
Suppressed Andreecium or Coane Sate: (Akiw cs. gene er OS 
Alone with suppressed Pertanthy <5 1. «os ste « = 194 
INetira eNO WeCsiu. saath ee commerce cM sls) oso as, ae ante a OD 

§ 6. INTERRUPTION OF NORMAL ALTERNATION . ... . . 195 
Anteposition or Superposition. . . .... ome Ca mee a eie 
eAippaarance only” 285 an Pea ie a ot Be PT 196 
Superposition by Spirals ... . Eanes ap LOG 
Anteposition with Isostemony and Ds eee aelomrepe Ws LOG 
MiEh@bdiplostemony ys). 2. sh Sacete 6. fe oe) OR eae tee 98 

Si) INCREASED NUMBER OF PARTS’. «29. 5 2%. » « « 200 
Remar vidlimplicaione 6 «she Ge) 4 ep ee ae on 200 
iParapetalous: Multiplication <9. oi ¢9 st aes ts ee 2) OL 
@horisissorMeduplication.. 2s) - = wares ot a 6202 

SSamOULGROWDHS 0 ses ele eso eee, St (eS RM SIE A. ‘DOO 
Their relation to Chorisis: -Trichomes. . . .... . . 209 
ComonmiomCrowir i203 ces, Bikes soon es aoe ae. . 210 
ipl ewe eaes (sec! coe Wea oe ee re ee eles ZIT 

§ 9. Forms or THE Torus OR RECEPTACLE. . «. ... . 2i1 
Stipe, ea Cae SEER ORES: CHER re One Pe are 1 
DISkar rere! : Se Fe 5; 
yp am COMOI Nes sateiy a po. Boa. ee ee or ictikrs eon ot LA: 

Section IV. ApapraTIoNs OF THE FLOWER TO THE ACT OF 
HGRIEGI TA MMTOND casein) ig ttl ft oem ae 2) Lees pete is 2A 
Reem eLNN GrENTURVAE 0 nctiecy ius! Bg") s<s--y iota” Rise at uk Waceeieele: .< 2L5 


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 


Irregularitv as related to Allogamy ......... 219 


CONTENTS. 


Dichogamy, either Proterandrous or Proterogynous . 
Proterogyny . 

Proterandry 

Particular Adaptanent in Pea pionaeeoe Flowe ers 

In Kalmia-blossoms, Iris, &c. 

Transportation of Pollinia . 

In Orchidacew and Asclepiadacee 

Heterogonous Dieter and menses 


§ 38. ADAPTATIONS FOR CLOSE FERTILIZATION . 


Cleistogamy . 


Section V. THE PERIANTH,. OR THE CALYX AND COROLLA IN 


PARTICULAR 


Perianth as to Duration, Numerical Terms, Union, &c. 
Parts of Petals and of Gamophyllous Perianth . 
Forms of Corolla and Calyx ‘ 


Section VI. Tue ANDR@CIUM, OR STAMENS IN PARTICULAR . 


The Stamen as a whole; Numerical Terms . ; 
The Filament and the Anther; their Modifications 
Pollen 

Pollen-tubes . 


Section VII. Tue Pistits, or GyN@&CIUM 
-§1. IN ANGIOSPERMS . 


Carpel or Carpophyll . oe At 

Ventral and Dorsal Sutures; Placenta 

Simple or Apocarpous Pistils . 

Compound or Synearpous Pistil . 

With two or more Cells and Axile Piacantee Partitions 
With one Cell and Parietal Placente 

With one Cell and Free Central Placenta 

Anomalous Elacentatiom << "= Welerss Were! bee: ae 


§ 2. In GYMNOSPERMS . 


Structure in‘Gnetacee . . . = «. =» « « 
Simin Commies es 5 9 6 6 o o - o 9 O-¢ 
In the Yew Family 

In the Pine Tribe, &e. 

In the Cypress Tribe . 

Structure in Cycadacee . 


Section VIII. Tue OvuLe 


Its Structure and Position . 


Its Forms, Orthotropous, Campylotropous, ee 


Anatropous 
Origin and Morgholamenl aan ae pas Omles 
Origination of the Embryo . 


CONTENTS. 


CHAPTER VII. THE FRUIT 


Secrion I. Irs Srructure, TRANSFORMATIONS, AND DEHIs- 
CENCE. 


Pericarp, its Alterations, Accessions, and Transformations 
Dehiscence 


Section II. Tue Kinps or Fruit 


Simple Fruits : jth 

Dehiscent Fruits, Follicle, Tenn. eaeaule iy xis Signe 

Indehiscent Dry Fruits, Samara, Akene, oe Caryopsis, 
Nut, &e. . . 

Fleshy Fruits, Drupe, pea pene ery. &e. 

Aggregate Fruits Se heh 

Accessory or Anthocarpous ineiies 

Multiple or Collective Fruits, Syconium, Strobile. &e. 

Table of Simple Fruits 


‘CHAPTER VIII. THE SERED 


Its Stalk, Coats, and ae 
Aril or Arillus : 

Nucleus or Kernel, Albumen 

The Embryo, its Parts and Positions 

The Cotyledons as to Adjustment and Sen: 


CHAPTER IX. TAXONOMY . 


Section I. THe PRINCIPLES OF CLASSIFICATION IN NATURAL 
History 


Individuals 

Species . 

Varieties, iaaece: digs 

Cross-breeds and Hybrids 

Genera . ; : 

Orders, Classes, Tribes, Ber 

Sequence of the Grades . one 

Nature and Meaning of Affinity . i 
Theory of Descent and Natural Selection 


Section II. Boranicat CLassiFICATION 


Ante-Linnzan Classifications . 
Linnean Classification 

Sexual Artificial bee 
Natural System 

As presented by esi ; 
Some of its Modifications 


xi CONTENTS. 


CHAPTER: X, PHYTOGRAPEY 5 cic. eee cyt eee 
Section 1. NOMENCLARURE” .4.)5 <5 *. 12) «0» 5h see eS 
Names of Plants, Binomial Nomenclature ..... . . 3846 
Rules fornaming:; Plants “is... = i cy cyte eee 
Names of Genera . . . PO ce ane errs 
Names of Species, Neeretien! on hotene ee en 50) 
The Fixation, Precision, and Citation of Gavies aE 
Subgeneric Names 2 5.9". 7 4 2) 58 
Tribal'and Ordinal’Names'. ~ 2 2% 4° .° 20a) 2 cee 
Names of Cohorts@lasses. Ge 35:5). ee EES 
Section II. GLossoLoGY OR TERMINOLOGY .-..... «. ooo 
Secrion JIL. DrscRIPTION.© : . 2956 6 6) 2 6 eon 
@haracters- 4.03. Ss oe ee ee, ne Ca 
Punctuation . "4 6 ee ee eer Se 
Synonomy «2 o eee oe ee 
Iconography. . . . © Be ee, G os 2a noe 
Habitat and Station, ee alee Dat PRL al er, ec 
Rtiymology of Names: =. 297 2) =) cue che cup Cc eerememmneLEee 
Accentuation, Abbreviations 4 .: . « =. . «| =» 0 thenneOOM 
Signs .. =. ee wr) we S| chat Sites aon um 
Floras, Marcie aoe ee res St 
Section IV. Specimens, D1irRECTIONS FOR THEIR EXAMINA- 
TION, PRESERVATION, GG) 9: « ‘:\ 2 | & [a = eeeomemCMD 
Implements of Investigation . . . . =. 5 = = |) sesneeene 
Diagrams...) 32+ -<p .b eof os ws ce gs 
Wer bOrizing) ss capers ss i 0 ee ge yeel eee et eee 
Drying Specimens.) 9. «cK ee 
Poisoning Specimens... 2. so. i ok se oe 
The Herbarium... ofS 2%. 266 oe 2S <b eee 
ABBREVIATIONS “he 6 st cs eae cee oe oy Ge San SCD 
SIGNS. 2.20 )..2 2 4.2.3 904-5 © 204 7 See 


GLOSSARY OF BOTANICAL TERMS, WITH INDEX . . 393 


STRUCTURAL BOTANY 


THE BASIS OF MORPHOLOGY. 


INTRODUCTION. 


1. Tue two Biological Sciences,’ considered as parts of Natural 
History, are Zoology 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, 7. e. of the two organic kingdoms, including 
both their physiology and descriptive natural history. 

1 


2 INTRODUCTION. 


of plants and animais are composed, is called organic matter. 
Animals appropriate and live upon this, but have not the power 
of producing it. ‘So the vegetable kingdom stands between the 
mineral and the animal; and its function is to convert materials 
of the one into food for the other. Although plants alone are 
capable of building up living structure out of mineral mate- 
rials, and are the sole producers of the organic matter which 
is essential to animal life, and although animals consume that 
which plants produce, yet 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 
assimilate mineral matter. Most plants of the higher grades 
assimilate largely and consume little, except in special opera- 
tions. Some, 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 Puysio- 
LOGICAL or BroLocicaL Borany (using these terms in their widest 
sense) and Systematic Botany. But as Physiology and Biology, 
in the restricted sense, relate only to functions or actions and 
their consequences, the first department naturally divides into 
two, viz. Structural Botany and Physiology. 

5. SrructuraAL Botany comprehends all inquiries into the 
structure, the parts, and the organic composition of vegetables. 
This is termed OrGANOGRAPHY, when it considers the organs or 
obvious parts of which plants are made up, and MorpHo.oey, 
when the study proceeds on the idea of type. The term 
OrGANOGENY has been applied to the study of the nascent 
organs and their development; Puyroromy, or VEGETABLE 
Anatomy, to that of the minute structure of vegetables as re- 
vealed by the microscope, 7. €. to the composition of the organs 
themselves. But, 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 Hisrotocy. The study of 
functions, or of the living being (animal or plant) in action, 
is the province of PuysroLocy. 

6. Systematic Borany, 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 Piants, the actual arrangement of known plants in 
systematic order according to their relationships ; PHyroGrapny, 
the rules and methods of describing plants ; and NoMENCLATURE, 
the methods and rules adopted for the formation of botanical 
names. GuLossoLocy or TrerMrINoLocy’ is a necessary 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.’ 

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 
physiology. But these will here be treated only in the most 
general or incidental and elementary way, 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, 7.e. into PH2xNoGaMmous (or PHANEROGAMOUS) or 
FLOWERING, and CryproGamous or FLoweriess Piants. The 
first are all manifestly of one type. 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 Grossotocy is the better word, but Terminoxoey, although a hybrid 
of Latin and Greek, is in common use. 

2 What is called GeocrapuicaL 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. Fossrr Borany (Vegetable Paleontology) relates to the 
plants of former ages, as more or less made known in their fossil remains. 
Mepicat Borany, AcricuLtuRAL Borany, 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 Phxnogamous 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, E 


OUTLINES OF THE GENERAL MORPHOLOGY OF 
PHA NOGAMOUS PLANTS. 


10. Morrpnoroey, 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 elossology and organography. 
But it deals with these from a peculiar point of view, and under 
the idea of unity of plan or type.? 

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 Geethe, at least as 
early as the year 1817 (Zur Naturwissenschaft iiberhaupt, besonders zur 
Morphologie, Stuttgart und Tiibingen, 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, A/etamorphosis. (304.) 

Apparently the first botanist to adopt the term was Auguste de St. 
Hilaire, in his “Lecons de Botanique, comprenant principalement la Mor- 
phologie Végétale, etc., Paris, 1841. The term seems not to have been taken 
up, in zoology, by Etienne Geoffroy 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 


and function. ‘These appendages, whatever their form or use, 
accord with leaves in mode of origin, position, and arrangement 
on the axis or stem. Their most general and ordinary form is 
the familiar one of foliage ; hence the name of leaves has been 
by botanists extended in a generic way from the green expan- 
sions which constitute foliage to other forms under which such 
appendages occur. The proper morphological expression is, 
that the latter are homologous with leaves, or are the homologues 
of leaves. 

13. Leaves are borne upon the stem at definite places, which 
are termed Nopes. 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 the cireum- 
ference of the stem equally, forming a circle, technically a 
Wouort, or in Latin form a Verticit. 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 are alter- 
nate, that is, come one after another on the stem; or in whorls 
(whorled, verticillate), in the commoner case of a single pair 
being called opposite. The bare space between two successive 
nodes is an IntERNOpDE. This is longer or shorter, according to 
the amount of longitudinal growth, which thus spaces the leaves, 
or, whorls of leaves, 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, 7. e. 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 Axms 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 stem, connected with the ground, produces 
roots which develop downwardly into the soil, from which they 
draw sustenance. Branches, when developed above ground, 


1 A common designation for all these appendages being desirable, a good 
one is furnished by the Greek name for leaf, @vAAov, PHyLiuM, plural 
Pura. This, used with prefixes, may be made to designate the kind of 
leaves in many cases, —as, prophylla, cataphylla, hypsophylla. 

Recent German botanists use the word Phyl/ome in this sense. It is a 
rather convenient and well-sounding word; but phyl/oma is the exact Greek 
equivalent of our word foliage, and therefore not very well chosen as a 
common term for leaves which are not foliage as well as those which are. 
Nor will this word, like phy//um, readily take prefixes, as above, or the adjec- 
tive form, as it readily does in prophyllous, hypsophyllous, gamophyllous, &e. 


OF PHASNOGAMOUS PLANTS. T 
being in organic connection with their parent stem, do not 
usually produce roots; but when placed in equally favorable 
conditions for it, 7. 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, 
isa Bup. 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.* 

16. As branches are repetitions and in one sense progeny of 
the stem which bears them, so the serial similar parts or leaf- 


bearing portions of a simple stem are repeti- Ne 2e yes 
tions, or in a like sense progeny, each of the VY By 
preceding one from which it grew. The g YW 
simple-stemmed plant is made up of a series 


of such growths, each from the summit of r 
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 Puytoxs. 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 PuyrOmera (qgetor, plant, | 
usgoos, 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 PH¥romEr. 
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 phytomera, 
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 Pheno- 
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 which the normal development of the axis finally 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 which sexual reproduction takes place, 
may be successively treated in this order. 


OF THE EMBRYO AND SEEDLING. i) 


CHAPTER II. 


MORPHOLOGY AND DEVELOPMENT OF THE EMBRYO AND 
SEEDLING. 


19. The Embryo is the initial plant, originated in the seed.? 
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. 5 
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 superfcetation in 
some seeds. In those of the cultivated Orange it is most common, and an 
evident monstrosity. -In Conifer and Loranthaceex, 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 


somewhat loosened. 3. Embryo of same, just beginning to unfold in germination. 
4. Same more advanced: a. its stem or caulicle; 0d. its two leaves o» cotyledons. 


10 MORPHOLOGY 


20. Caulicle or Radicle, and Cotyledons. The name of radicle 
was early applied to the axis of the 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 (24, 
44, 78), but a body of the exact nature of stem, from the 
naked end of which the root is developed. Wherefore Caulicle 
(Lat. cauliculus, diminutive of caulis, stem) is 
the appropriate name; and it would be gen- 
erally adopted, were it not that the older term 
is so incorporated into the language of sys- 
tematic botany (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 or%an, is preferable.t The more fanciful 
name of Cotylédons 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, 7. 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- 
i H ling instead of coiling of the cotyledons in the 
' 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 here be described, taking that of 
a Maple for a convenient type or pattern, with which other forms 


1 Linneus called it Rostellum, a name which, being etymologically mean- 
ingless in this connection, is not misleading. The French botanists named it 
Tigelle, diminutive of tige, stem: but some (like Mirbel) applied the term to 
the developing axis above the cotyledons; others, to the early axis both 
above and below them. The name Radicula originated with Gaertner. 

2 The name Cotyledon, which was adopted by Linneus, is a Greek word 
for a cup-shaped hollow or cavity, also for a plant with thickish and saucer- 
shaped leaves. It was primarily applied to the thickened “lobes” of the 
embryo, the foliaceous nature of which was not recognized. 


FIG. 5. One of the twin winged fruits of Red Maple (Acer rubrum), with body 
divided, to show the seed. 6. Seed extracted and divided, to show the embryo within. 
vt. Embryo partly unfolded. 8. Embryo in early stage of germination. 


OF THE EMBRYO AND SEEDLING. Fk 


may 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 they 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 they are called, 
7.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 
Tudiments of the second: the lower portion covered with root-airs is the root; the 
naked portion above is the canlicle. 


12 MORPHOLOGY 


After this is consumed and in good part converted into struc- 
ture, the plantlet must by the action of 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. 9), next as somewhat upraised 
by the development of the second internode 
(Fig. 10, summit), and finally both this inter- 
node 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 internode 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. But, 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.’ It is formed 


ul 


1 Yet from nothing which is special to this part of the embryo, nor to the 
embryo at all. The primary root is developed from subjacent tissue of the 
tip of the caulicle, just as it is sometimes developed from along the sides, 
and as secondary roots are from all or most stems under favoring conditions. 
This complete similarity, and the fact of what is called the “endogenous ” 
origin of roots (i.e. their springing from subjacent rather than superficial 
tissue) appear fully to warrant the statement in the text above. 


FIG. 10. Maple plantlet with second internode developing. 11. Same with second 
internode and pair of leaves complete, and bud of the third apparent. 


OF THE EMBRYO AND SEEDLING. a kes) 


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 


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 distine- 
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 they grow either leaves 
or any organs homologous with leaves. 
They 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, Ipomea purpurea, or any 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 


12 


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 plantlet, until that has established itself 
and had time to elaborate proper material therefor. This con- 
dition is correlated with thin foliaceous 
A BS cotyledons, holding no store of nourish- 
*/ ment. Here they do not contain sufficient 
material for the development of the initial 
stem and root. The maternal provision 
for this is here stored up in the seed 
around but not within the embryo. This 
nourishing deposit, seen in the section 
(Fig. 15) filling the whole space between 
the seed-coats and the thin embryo, was 
named by the early botanists and vege- 
table anatomists the ALBUMEN of the seed.* 
This substance, softened in..germination 
and by chemical changes rendered soluble, 
is gradually absorbed by the cotyledons 
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 exal- 
buminous, those supplied with and those 
destitute of albumen. The difference 
inheres neither in the character nor in 
the amount of the maternal provision for the development of 
the embryo-plant, but merely in the storage. In exalbuminous 
seeds the nourishment supplied for this purpose is taken into 
the embryo itself, mostly into the cotyledons, during the growth 
and before the maturity of the seed. In albuminous seeds 
this same material is deposited around or at least external to 
the embryo. 

27. The amount of this deposit is, in the main, inversely pro- 


1 Grew appears to have first applied this name, and Gaertner to have 
introduced it into systematic botany, where it remains in use, although 
Jussieu replaced it by the term Perisperm, and Richard by Endosperm, 
neither of them much better etymologically than the old word Albumen. 
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 egg as a body or mass, and not as 
a chemical substance; the embryo being fancifully conceived to be analo- 
gous to the yolk of the egg, the surrounding substance of this kind not 
unnaturally took the name of the white, viz. albumen. 

FIG. 13. Section of seed of common Morning Glory, Ipomcea purpurea, dividing 
the contained embryo through the centre. 14. Embryo of same, detached and straight- 
ened. 15. Embryo in germination; the cotyledons only partly detached from the coat 
of the seed. 16. Same, later and more developed, the cotyledons unfolded and out- 
spread as the first pair of leaves. 


OF THE EMBRYO AND SEEDLING. 15 


portional to the size and strength of the embryo, 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 embryo left in 


2 al 19 7 


™ 


white ; the albumen 3 


24 22 20 
as a dotted surface. 


The lower rank shows the einbryos 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 in 
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- 
bryos there is no apparent distinction of parts until they develop 
in germination. 

28. The study of the formation of the embryo 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 embryo 
while yet in the seed. It also appears that the size and shape 


FIG. 17. Section of seed and contained embryo of Mirabilis (IFour-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 organ do not indicate its nature, either in the embryo or. 
in subsequent growth. But in all 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 
different 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, 


8 cre 


or a primary terminal bud, is visible in the seed. It is most 
manifest in large and strong embryos with thick or fleshy cotyle- 


FIG. 25. Further development of Morning Glory, Fig. 16, the root cut away, the 
internode above the cotyledons and its leaf completed, the next internode and its leaf 
appearing. 

FIG. 26. Embryo (kernel) of the Almond. 27. Same, with one cotyledon removed, 
to show the plumule, a. 

FIG. 28. Section of an Apple-seed, magnified, cutting through the thickness of the 
cotyledons. 29. Embryo of the same, extracted entire, the cotyledons a little separated. 

FIG. 30. Germination of the Cherry, showing the thick cotyledons little altered, 
and the plumule developing the earliest real foliage. 


OF THE EMBRYO AND SEEDLING. LT 


dons, ?. 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 itis 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 cherry, 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 (2.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- 
bryos 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; they 
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, 
haying 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 internode and a pair of leaves. 

2 


18 MORPHOLOGY 


latter are practically the earliest efficient foliage of the plantlet. 
Thus, as in the germinating Cherry-seed (Fig. 30), three or four 
internodes of stem, with their leaves, may be produced before 
these leaves themselves are sufficiently developed to make any 
sensible contribution to this growth. And in the Beech and Bean, 


the leaves of the plumule come forward almost before the root 
has attached the plantlet to the soil. (Fig. 32, 35.) Between 
such cases and that of Maple and the like there are all degrees. 
There are also familiar cases in which the storage of nourishment 
in the cotyledons is carried to a maximum, with results which 
gravely affect the development. 


FIG. 34. The embryo (the whole kernel) of the Bean. 35. Same early in germi- 
nation; the thick cotyledons expanding and showing the plumule. 36. Same, more 
advanced in germination; the plumule developed into an internode of stem bearing a 
pair of leaves. 

FIG. 37. Embryo of Pea, i. e. 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 
relative 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 sime 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 


40 


buried beneath the surface of the ground, there it remains. The 
abortion of the earliest leaves of the plumule is in correlation 
with this Aypogeous (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. 
the same. 


40. Advanced germination ot 


20 MORPHOLOGY 


33. In some Oaks, notably in our Live Oak (Quercus virens), 
and less so in the Horsechestnut, the two cotyledons coalesce or 
cohere by their contiguous faces. 
In some of these cases of hypo- 
geous germination, the short 
ceaulicle and plumule are extri- 
cated from the enclosing coats or 
husk by the development of short 
stalks (petioles, 157) to the fleshy 
cotyledons ; as is seen in Fig. 42, 
and in most germinating acorns. 
These petioles are not visible in 
the seed, but are 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 
formation of far longer stalks 
(petioles) than those of the 
Horsechestnut. A singularly dis- 
guised instance of this kind is seen in Megarrhiza, a genus of 
Cucurbitaceous 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 off 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, the 
state of things represented in Fig. 43 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 cleft 
below. This explains the anomaly. The real caulicle has re- 


FIG. 41. Section of a Horsechestnut or Buckeye seed. through the very thick 
cotyledons and the incurved caulicle. 42. Seed in germination, showing the petioles 
to the cotyledons, &c. 


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 
Cucurbitaceze to 
bring up the cotyle- 
dons that they may 
develop as leaves 
(as in the Pumpkin, 
Fig. 47): here this 
elevation is brought ut 43 
about in a different way, but without securing the useful end.’ 


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. 


a 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 species of Larkspur (notably in the Californian Delphin- 
ium nudicaule) ; but in these the cotyledons develop into a pair 
of efficient green leaves. 


if 


45 
36. A similar elongation of petioles of the cotyledons, but 
without any union, occurs in a species of Morning Glory of the 
plains beyond the Mississippi (Ipomeea leptophylla) ; 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 (25, Fig. 15, 16, 25), and note that the difference is merely 
that the caulicle in the common Morning Glory 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 caulicle), and forms such a direct downward 


FIG. 45. Germination of Ipomca leptophylla; the caulicle not developing, the 
plumule and the petioled cotyledons rise from underground. Dotted line marks the 
level of the soil. 

FIG. 46. Embryo of a Pumpkin, the cotyledons separated. 47. Same germinated ; 
a cluster of roots from the base of caulicle. 


OF THE EMBRYO AND SEEDLING. 23 


prolongation of it, that it was called the descending axis; and 
the body 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; 7. e., it strikes root in 
nearly the mainer 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 polycotyledonous | 
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 Coniferze, a whorl of from 3 to 10 
cotyledons isshe normal structure, varying accord- , 
ing to the species, but of almost uniform number | 
in oh. 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 K 
into a circle of needle-shaped leaves. In the Pine * 49 
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; 
z. e., are either opposite or verticillate. From the occasional 
union at base of the cotyledons of a polycotyledonous embryo in 
pairs or groups, and from a study of their early development, 
Duchartre’ plausibly maintains that such cotyledons really consist 
of a single pair, parted into divisions or lobes. The ordinary 
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. 


24 MORPHOLOGY 


of the plumule; that is, the leaves of the embryo are alternate. 
But the caulicle is usually very short, and there is no external 
mark by which its limits may be distin- 
guished from the cotyledon, until germi- 
nation has begun. For a type of it, the 
embryo of some aquatic or marsh plants 
may be taken, where it forms the whole 
S A kernel of the seed (Fig. 50-53), and 
50 51 52 53 the structure can be made out antecedent 
to germination. It is understood by 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 conecreted, except a minute 
longitudinal chink at base, out of which the growing 
plumule protrudes in germination. The embryo of 
Iris may be similar in structure, but no distinction 
of parts is visible. 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 successively 
enclosing the inner and younger. (Fig. 55.) Here, 
therefore, the cotyledon mainly remains in the seed, 
and the seed remains underground (hypogzeous). 
40. It is somewhat different 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 first steps are the 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 
filiform green leaf, which, taking an erect position, 


FIG. 50. Seed of Triglochin palustre; the rhaphe, leading to the strong chalaza at the 
summit, turned towards the eye. 51. The embryo detached from the seed-coats, showing 
the longitudinal chink at the base of the cotyledon; the short part below is the radicle. 
52. Same, with the chink turned laterally, and half the cotyledon cut away, bringing to 
view the plumule concealed within. 53. A cross-section through the plumule, more 
magnified. Q 

FIG. 54. Section of seed of Iris, enlarged, showing the small and apparently simple 
embryo at the base of the albumen 55. Germinating seed and seedling of the same, of 
natural size. 


OF THE EMBRYO AND SKEDLING. 25 


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 internodes remain undeveloped until the 
plant is ready to blossom. Very similar is the germination of 
a date-seed, except that the, 2 y} @, HLS 
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 iT 
and the manner in which «-~ 
its rudimentary leaves 3. | 
successively enclose gr FAN 
other. That of Maize or, 
Indian Corn, one of the \W4 
largest, is most convenient §© ° Gop s 4 
for study. (Fig. 56-59.) 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 cnwraps the plumule: it adheres 
closely to the albumen by the whole back, and remains un- 
changed in germination: its function is to absorb nutritive 


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 dotted lines. 

FIG. 59. Vertical section of Indian Corn across the thickness of the grain, dividing 
the embryo through the centre and displaying its parts: ¢, 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 MORPHOLOGY 


matter furnished by the albumen, and to transmit it to the. 
growing plumule. ythe, giueues consists of a succession of 
vudimentary leaves, sheathing 
. and enclosing one another, on 
the summit of a very short 
-axis, which is mainly the 
‘caulicle, otherwise called rad- 
jicle. This is completely en- 
closed by a basal portion of 
the cotyledon and of the 
-outermost leaf of the plu- 
mie, which form a peculiar 
‘sheath for it, named the 
“Coleorhiza,* i. e. 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. 62, 
J 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 Pseudo-monocotyledonous 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, 18, 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 


i\ 


2 


1 This, the Coleorhize of Mirbel, should not be confounded (as by some it 
has been) with the “root-cap,” or tissue which ordinary roots (whether 
primary or secondary) break through in their development or carry on 
their apex. 


FIG. 62. Early germination of Indian Corn. . More advanced germination ot 
same: roots produced from portion of stem above in ‘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 Phznogamous plants, viz. DicoryLepongs or DicoTyLEDONOUS 
Priants, and MonocotyLepones or MonocoryLeponous Pants, 
— names introduced into classification by Ray, and adopted by 
A. L. Jussieu, in his Genera Plantarum. 


CHAP TER - Ui: 


MORPHOLOGY AND STRUCTURE OF THE ORGANS OF THE 
PLANT IN VEGETATION. 


Section I. Or tHe Roor. 


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 may elaborate into nourishment. As 
already 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 may 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; 5. the older and firmer tip. 


28 MORPHOLOGY OF THE ROOT. 


is similarly capped or protected.’ But the so-called root-cap is 
seldom so distinct or separable as to deserve a particular name. 

45. Nature of Growth, 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 type, that they take one common name, that of CELLs. 
These are the histological elements of plants, 7. e. the units of 
minute anatomical structure. While, in the morphology of the 
plant’s obvious organs, analysis brings us to the phytomer (16) 
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 (cellula, by the French 
conveniently termed cellule) is the 
living vegetable unit, in the same 
sense that the brick is the unit 
of a brick edifice. 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 the 
65 } forming structure by a kind of 
«6 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 growth.2 The inspection through a 


1 The notion that the tip of the root consists of delicate forming or 
newly formed tissue, or bears some organ or structure of this nature (a 
“ Spongiole”), has hardly yet been eliminated from the text-books and popular 
writings. It had no proper foundation in fact. 

In Lemna, and in some other aquatics, and also in some aerial roots, this 
older tissue often separates into a real root-cap, free at base, like an inverted 
calyptra. 

2 This, as to the structure, is the subject of Histology; as to processes or 
actions, the subject of Physiology ; both to be treated in a separate volume. 

FIG, 65, 66. Portions of surface of Fig. 64, more magnified, clearly displaying the 
superficial cellular structure and the long processes from some of the cells, called root- 
hairs, whieh abound on the upper part of Fig. 64. 


MORPHOLOGY OF THE ROOT. 29 


simple microscope of a slender young root, and of thin slices 
of it immersed in water, may serve to give a general though 
crude idea of the vegetable cellular structure, sufiicient for the 
present purpose. Roots are naked; that is, they bear no other 
organs. When they send off 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 are 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. (3583.) 

46. 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 young roots increased by the 
production of root-hairs. (Fig. 64, upper part, and more magni- 
fied in Fig. 65, 66.) 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 Maclura or Osage Orange so readily originate 
buds that the tree is commonly propagated by root-cuttings. 

48. Kinds of Roots. The root, commonly single, which origi- 
nates from the embryo itself, is called the Prmotary Root. (37.) 
Roots which originate from other and later parts of the stem, 
or elsewhere, are distinguished as SeconpAry Roots. But the 


30 MORPHOLOGY OF THE ROOT. 


latter are 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, &c. It is mainly annuals and common trees that 
naturally depend on the primary root; and most of these can be 
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, cither 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 weli marked. 

00. Annuals are plants which, springing from the seed, flower 
and seed the same year or season, and die at or before its close. 
They produce fibrous roots, either directly from the embryo and 
succeeding joints of stem (as in Grasses, Fig. 63), or from a 
persistent primary or /ap-root, more or less thickened into a trunk’ 
or divided into branches. The products of vegetation in all such 
herbs are not stored in subterrancan 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. 

51. 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 with the original root 
have 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 belong, become annuals 
in temperate climates, early perishing by autumnal cold. 

52. 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 rainy 
winter and rainless summer, many germinate in autumn, vegetate 


MORPHOLOGY OF THE ROOT. 31 


through the winter, flower and seed in spring, and perish in 
early summer. These may be termed WryTER 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.? 
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 and 
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. 


32 MORPHOLOGY OF THE ROOT. 


54. But some plants, such as the Radish, which when they 
spring from seed in autumn are true biennials, will when raised 
in spring pass on directly to the flowering stage in summer, or 
when sown after the warm season begins will often run through 
their course as annuals. Then there are various biennials which 
thicken the root very little and hold their leaves through the 
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. 

55. 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 fibrous roots, 
these are produced anew from time 
to time or from year to year. Also, 
while 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. 68) and the Peony afford good 
examples of this. Sweet potato is - 
another instance.t Most such roots 
have only a biennial duration: they 
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, corms, 
bulbs, &c. (See 115-122. 

56. The distinction between annuals and biennials is at times 
so difficult, 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 DeCandolle to unite 


1 It is only by the readiness of this root to produce adventitious buds, 
especially from its upper part, that it has been mistaken for a tuber, such 
as the common potato. 


FIG. 68. Fascicled and tuberous or fusiform (secondary) roots of Dahlia: a,a. buds 
on base of the stem 


MORPHOLOGY OF THE ROOT. 33 


the two under the common appellation of Monocarpric plants, 
Plante monocarpice, taken in the sense of only once-fruiting 
plants; and to designate perennials by the corresponding term 
of Potycarpic, Plante polycarpice, literally many-fruited, taken 
in the sense of many-times fruiting.’ 

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 they 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 ba8e, 
small offsets to blossom the next season; and so on. Then 
Houseleeks (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 years ; 
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 yegetates 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. Mondtocous (bearing progeny once) and Polytocous (bearing 
many times) would be more appropriate. 

3 


34 MORPHOLOGY OF THE ROO?Y. 


into which they penetrate. More remarkable cases abound in 
those tropical regions where the sultry air, saturated with moist- 
ure for a large part of the year, favors the utmost luxuriance of 
vegetation. In the Palm-like Pandanus or Screw-Pine? (Fig. 
69), very strong roots, emitted in the open air from the trunk, 
and soon reaching 
the soil, give the 
appearance of a tree 
partially raised out 
of the ground. The 
famous Banyan-tree 
of India (Fig. 71) is 
a still more striking 
illustration ; for the 
aerial roots strike 
from the horizontal 
branches of the tree, 
often at a_ great 
height, at first swing- 
ing free in the air, 
but finally 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 
the Pandanus, but also from the branchlets, as in the Banyan. 
Even the radicle of the embryo starts into growth, protrudes, 
and attains considerable length while the fruit is still attached to 
the branch. 

59%. Aerial Rootlets for climbing are familiar in the Ivy of the 
Old World (Hedera), Trumpet-Creeper (Tecoma radicans), and 
our Poison Ivy (Rhus Toxicodendron) ; by the adhesion of 


1 So named, not from any resemblance to a Pine-tree, but from a like- 
ness of the foliage to that of a Pine-Apple. 


FIG. 69. Pandanus, or Screw-Pine; and in the background, 70, a Mangrove-tree 
(Rhizophora Mangle). 


MORPHOLOGY OF THE ROOT. 30 
which the stems, as they grow, ascend walls and the trunks of 
trees with facility. In Rhus a superabundance of these rootlets 
is produced, thickly covering all sides of the stem. 


60. Epiphytes or Air-Plants also have roots which are 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 phznogamous plants, are in this case. The greater 
part of the phenogamous Epiphytes pertain to two monocotyle- 
donous orders, the Orchis family and that to which the Pine- 
Apple belongs, viz. the Bromeliaceze. 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, 7. e. plants of the Orchis family, are the 
most showy and numerous, and of the greatest variety of forms, 
especially of their blossoms, which are often bizarre and fantas- 
tic. They belong, naturally, to climates which are both warm 
and humid: they 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 visit them. In some the blossoms curiously 


FIG. 71. The Banyan-tree, or Indian Fig (Ficus Indica). 


36 MORPHOLOGY OF THE ROOT. 


resemble butterflies or other insects; as, for example, Oncidium 
Papilio, Fig. 72. Epiphytic orchids are indigenous to the United 
States only from Georgia to Texas, and only in humble forms, 
in company with species of Tillandsia, representing Bromeliace~ 
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 gray 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 fix themselves to other plants, 
but draw therefrom their nourishment, at least in part. Among 
cryptogamous plants very many Fungi are parasitic upon or 
within living plants or animals. But only pheenogamous para- 
sites are here under consideration. These may be divided into 
two classes ; those with and those without green foliage. 

62. Green Parasites may be either wholly or partially parasitic ; 
that is, they may draw all their support from a foster plant, or 


FIG. 72. Oncidium Papilio, and, 73, Comparettia rosea; two showy epiphytes of the 
Orchis family; showing the mode in which these 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, they 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 by 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 £7 
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 


\\ 


PY 
‘i 
¥ 


iss 


FIG. 74. Native epiphytes of Georgia, &c.: the erect one at the right an Orchid, 
Epidendrum conopseum; the hanging one Tillandsia usneoides, called Long Moss. 

FIG. 75. Roots of Gerardia flava: some of the rootlets attaching themselves para- 
sitically to the root of a Blueberry. (From @ drawing by Mr. J. Stauffer.) 


3 MORPHOLOGY OF THE ROOT. 


America (called Cursed Fig) are examples of the former. They 
form trees, send down aerial roots in the manner of the Banyan ; 
but, while some roots seek the ground, some may attach them- 
selves to other trees parasitically, 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 Gerardia (at least of the section 
Dasystoma) and other plants of the 
same family, the uncultivability of 
Which is thereby explained. Also 
@) in Comandra and in their relatives 
‘the Thesiums of the Old World, 
belonging to a aie order (the Santalaceze) which has much 
affinity aie the entirely parasitic order (Loranthacex) to which 
the Mistleto belongs. 

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 discs, 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, 
2.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, 

7 78 7 it forms suckers, which attach 
themselves firmly to the surface of the supporting plant, 
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. Thus stealing its nour- 


FIG. 76. Section of one of the attached rootlets of Gerardia, showing the union. + 

FIG. 77. The common Dodder of the Northern States (Cuscuta Gronoyii), 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. 79. The same in germi- 
nation, elongating into a thread-like leafless stem. 


2 


MORPHOLOGY OF THE ROOT. 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.’ 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 they 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 stem 
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. Rafilesia 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 MORPHOLOGY OF BUDS. 


Section II. Or Bups. 


65. Buds are the germs of stems: they are axes with their 
appendages in an early state. Lrar-sups (GEMM) are those 
devoted to vegetation, and the parts, or some of them, develop 
as leaves. Mixep subs contain both foliage and flower or 
flowers. FLOWER-BUDS (ALABASTRA) are unexpanded blossoms. 
These are considered in another chapter. 

66. The conspicuous portion of an ordinary bud, or that which 
first develops, usually consists of leaves, or scales the homologues 
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 Houseleek (Fig. 91°), a Barberry 
and the Larch: when the internodes lengthen, the leaves are 
interspaced upon the axis. 

67. The cotyledons and plumule of the embryo are, morpho- 
logically, the first bud, on the summit of the initial stem, the 
caulicle. 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 
apical or 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 Jatent. 

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 
marked by the production of scales (Bud-scales, Perule, ete.) 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, Linnzeus gave to such bud-coyer- 
ings the common name of Hisernacutum. From the usually 
squamose (scale-like) character of this covering, such buds take 
the name of 

-69. Sealy Buds. Large and strong ones of this kind, such as 
those of Horsechestnut, Magnolia, Hickory, Lilac, &c., may be 
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 effectually, they are sometimes 
coated with a waxy, resinous, or balsamic exudation, as is con- 
spicuous on the scales of the Horsechestnut, 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-seales. That they answer to leaves is made 
manifest by « consideration of their 
situation and arrangement, which are 
the same as ofthe proper leaves of 
the species ; and by the gradual transi- 
tions from the former to the latter in 
many plants. In the 7urions, or sub- 
terranean budding shoots of numerous 
perennial herbs, and in the unfolding 
buds of the Lilac and Sweet Buckeye 
(4Esculus 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: 253), 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 the 
terminal bud; and below exhibiting the large rounded lJeaf-scars, as well as the rings 
or annular sears 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 


leaf-buds. 


MORPHOLOGY OF 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 commences.  Phila- 
delphus and Taxodium are of 
this kind. 

72. Subpetiolar Buds. Some 
leaf-buds are singularly cov- 
ered in their early state and 
through the summer, as in the 
Locust (Robinia), Honey-Lo- 
cust Fig. 96 (where they re- 
main very undeveloped), in 
Yellow Wood (Cladrastis). and 
more conspicuously in the 
Plane-tree (Platanus, Fig. 87): 
here they are all formed un- 
der the base of the protecting 
leaf-stalk, which in Plane-tree 
forms a sheath or inverted cup, 
very like a candle-extinguisher, 


fitted to and concealing the conical bud until autumn, when by 
the fall of the leaves these buds are exposed. 


FIG. 83. 


Diagram of vértical section of a strong bud, such as of Horsechestnut. 


84. The axis of the same developing, the elongation beginning with the lowest inter- 

node, soon followed by the others in succession. 85. A year’s growth of Horsechestnut, 

crowned with a terminal bud: a, scars left by the bud-scales of the previous year: ~ 
6, scars left by the fallen leaf-stalks: ec, axillary buds. 


FIG. 86. 
FIG. 87. 


Branch and buds (all axillary) of the Lilac. 
Leaf-bud under the petiole of the Plane-tree. 


MORPHOLOGY OF BUDS. 43 


73. Fleshy Buds. ulbs are peculiar buds of certain herba- 
ceous plants, with fleshy scales, and often of a more permanent 
character. Their nature and economy may most conveniently 
be illustrated under subsequent sections. Usually bulbs are 
subterranean or partly so. But small bulbs ( Bulblets, 123) regu- 
larly appear in the axil of nearly all the leaves of certain common 
Lilies, being obviously ordinary axillary buds, under certain 
modifications. They become detached at maturity, fall to the 
ground, produce roots, and grow as independent plants ; and 
their fleshy scales are storehouses of nourishment for the early 
support of this independent growth. 

74. Bud-propagation is a normal mode of reproduction in cases 
like the above, the- spontaneously detached bulblets or buds 
establishing themselves as progeny. In several species of 
Allium (Onions and Leeks), such bulblets usurp the place of 
flower-buds, making the analogy seem closer. Stems or branches 
which habitually root in the soil, or along its surface, equally 
propagate or divide into new individuals, becoming distinct by 
the perishing of the older connecting parts, or by breaking away 
from them. Propagation by cuttings is an acceleration or exten- 
sion of this same natural operation. The cutting is a portion of 
stem bearing one or more buds, which, through the faculty of 
the stem to strike root, is made to grow independently. In 
grafting, such a cutting, and in budding a bud only, with a small 
portion of wood and bark, is transferred to the stem of another 
plant of the same or of some related species, and made to grow 
there, uniting its wood and bark with those of the stock, and so 
becoming a limb or branch, in place of striking root into the soil 
and becoming a separate plant. The horticultural advantage of 
bud-propagation is, that the offsets or new individuals share 
in all the peculiarities of the parent as completely as if still 
branches of that tree. In propagation by seed, the special 
peculiarities or excellencies of individuals or varieties may not, 
and im some measure probably will not, be reproduced. 

75. Normal or Regular Buds, as to position, are either terminal 
or axillary, as already stated. (15.) They are single, that is, 
one bud normally occupies the apex of a stem or branch, and 
appears, or usually may appear, in the axil of (or upper angle 
formed with the stem by) any well-developed leaf. In these 
positions, buds are so usual, or so capable of appearing, that 
they are commonly regarded as potential when not actually 
present. The potentiality may be manifested by the actual 
development of these buds in shrubs or trees after the lapse of 
years. (84.) The terminal leaf-bud is to continue the axis it 


44 MORPHOLOGY OF BUDS. 


surmounts: axillary and any other lateral leaf-buds are to be- 
come branches. But even of buds which actually appear a large 
proportion do not grow. When a terminal bud is formed (as 
Fig. 81, 85, 91), this is commonly the strongest, or among the 
stronger. But in many cases it habitually or commonly fails to 
appear. In the Elm (with leaves and therefore buds 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. 86.) When all the 
regular buds make their appearance, and the leaves are opposite, 
the stem will be crowned with the terminal bud, having an axil- 
lary bud on each side of it. (Fig. 88.) 

76. Aecessory Buds. 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 (sperposed ), 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), &e., 
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, the 
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 Aristolochia Sipho, except 
that the uppermost bud is there strongest. 


FIG. 88. Branch of Red Maple, at the middle bearing triple axillary buds, placed 
side by side. 

FIG. 89. Piece of a branch of the Butternut, with accessory buds placed one above 
another: a, the leaf scar: 6, proper axillary bud: ec, d, accessory buds. 

FIG. 90. Part of a branch of Tartarean Honeysuckle, with crowded accessory buds 
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 accessory 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 ewtra-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 internodes 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 stems 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 II. Or tHe Srem. 
§ 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 phaznogamous plants possess stems. In 
those which, in botanical descriptions, are said to be acaulescent, 
or stem/ess, 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 
vegetation are chiefly carried on, not so much by the stem it- 
self as by a distinct 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. (13, 23.) 

80. Development and Structure. Ina 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, 
and under different conditions. Unlike 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 
may 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 deveiopment 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 
before growth begins, —also upon the amount of nourishment 
provided. Strong buds commonly have their parts, or some of 
them, ready formed in miniature, and a store of elaborated nour- 
ishment 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. 81, Horsechestnut, Fig. 85, and 
Hickory, Fig. 91) 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 GROWTH. 47 


the bud is well supplied with nourishment in spring by the stem 
on which it rests, 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 woody 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, may be traced back on the stem 
for a series of years, growing fainter with age, 
until they 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 6), the Magnolia (Fig. 81), 
and Hickory, Fig. 91. 

83. Ramification. Brancues (14-16) are secondary stems 
developed from a primary one, or tertiary ones from these, and 
soon. Ultimate or small ramifications of latest order are some- 
times called BrancuLets. The terminal bud continues the stem 
or axis which bears it. Lateral buds give rise to branches.’ 
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 phenogamous plants that it may here be left 
out of view. 

In phenogamous 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 TricuomEs (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 wiil be alternate; when the leaves are opposite, and 
the buds develop regularly, the branches will be opposite, &e. 
This holds in fact sufficiently to determine 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 ramification. 

84. Non-development of Buds. Someof 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 for 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 Horsechestnut (Fig. 85) ; sometimes the terminal one 
fails, and the lateral ones grow, when the stem annually becomes 
two-forked, as in the Lilac, Fig. 86. The undeveloped buds 
do not necessarily perish, but are 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, which the for- 
mer would have monopolized. 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 (76) 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. 96. 

86. Excurrent and Deliquescent Stems. 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 del/quescent, 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 rudimentally 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 
nnmatured, 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. Many of the details and applications of ramification, of 
most importance in morphology and descriptive botany, relate 


50 MORPHOLOGY OF STEMS. 


to anthotaxy or inflorescence (Chap. V.), which has its own 
terminology. But some of its terms may be conveniently 
employed in the description of ramification unconnected with 


flowering. 


§ 2. Forms or STEMS AND BRANCHES. 


89. On the size and duration of the stem the oldest and most 
obvious division of plants is founded, namely, into Herbs, 
Shrubs, and Trees. 

90. Herbs are plants in which the stem does not become 
woody and persistent, but dies annually or after flowering, down 
to the ground at least. The difference between annual, biennial, 
and perennial herbs has already been pointed out in the chapter 
on the root (50-57), and the gradations between them indicated. 
Herbs pass into shrubs and shrubs into trees through every gra- 
dation. The following definitions are therefore only general : — 

91. Undershrubs, or Suffruticose plants, are woody plants of 
humble stature, their stems rising little above the surface. If 
less decidedly woody, they are termed Suffrutescent. 

92. Shrubs are woody plants, with stems branched from or 
near the ground, and less than five times the height of a man. 
A shrub which approaches a tree in size, or imitates it in aspect, 
is said to be Arborescent. 

93. 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 single 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 still be called shrubs. 

94. 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 sufliciently 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 
(12) with stems, but not always. ‘Two kinds of erect stems 
have special names in descriptive botany. 

95. Culm is a name applied to the peculiar closed-jointed stem 
of Grasses and Sedges, whether herbaceous, as in most Grasses, 
or woody or arborescent, as in the Bamboo. 

96. Caudex is the name technically applied to the trunk of 
Palms (Fig. 126), Tree-Ferns, and the like, consisting of a 
commonly simple column, the surface beset with scales, — the 


SPECIAL FORMS. oh 


bases of former leaf-stalks, — or marked by scars, left by their 
fall. This name was used by botanists anterior to Linnzeus 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 stems of this sort, 
whether rising somewhat above or concealed beneath the surface 
of the soil. 

97. A Seape 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 
usually 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. Seandent or Climbing Stems 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.? 

100. Voluble or Twining Stems, 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 ‘‘ with the sun” (that is, from right to left of the 
observer viewing the coil from the outside”), 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 Dertrorse 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. 891) 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. Linneus supposed the observer to view the 


52 MORPHOLOGY OF STEMS. 


by coiling in the opposite direction, as the Bean (Phaseolus), 
the woody Aristolochia Sipho, the Morning Glory (Fig. 91*) and 

other Convolyu- 
lace. The Dod- 
der, a leafless par- 
asitic plant of the 
latter family, not only gains suppoit by coiling 
on the stems of other plants, but by attachment, 
through the development of sucker-like discs, 
along the whole contiguous surface. (Fig. 77.) 
The various actions through which plants climb, 
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 

ola 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, 
Tropzolum, and Solanum jasminoides (Fig. 255) ; 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 Cobzea 
and the Pea. 

102. Tendril-Climbers (Fig. 92-95) 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 stem up to the 


coil or circle from the inside; Mohl, Palm, Braun, and the DeCandolles 
adopt this, and the latter insist on it. Such authority should be decisive, 
if common usage and popular sense went along with it. But some of the 
botanists following Linneus adopted the reverse view; and to the present 
writer, as to Bentham and Hooker, Darwin, Eichler, and in part G. Henslow, 
it was so natural to view the coil from the outside that we without concert 
adopted this position and mode of expression. A right-hand coil, or one 
turning to the right, with us, is one the turns of which pass from the left to 
right of a bystander who confronts the coil. It is in this sense that a com- 
mon screw is called a right-handed screw, and that the right bank of a river 
is that to the right of the person who follows the course of the stream. So 
natural is this, that even on a map or plate, which has face and back, and 
therefore a right and left of its own, the figures occupying its right or left 
portions are understood to be those which are toward the right or the left 
hand of the observer who stands before it. 


FIG. 91%. Dextrorsely twining stem of Morning Glory, Ipomca purpurea. 


vet Te 
eo 


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 (597), 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. Stems 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 Assurgent, 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 (Sureulus) is an ascending stem rising from a 
subterranean creeping base. The Rose and Raspberry multiply 
freely by 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. 91") 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- 
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. 914. 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 branch 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. 

“410. 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. 92) ; 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 


FIG. 92. End of a shoot of the Grape-vine, with young tendrils: a sympodial 


stem. (See note.) 


FIG. 93. A portion of a stem of Ampelopsis quinquefolia, or Virginia Creeper, with 


a leaf and a tendril. 


FIG. 94. Ends of the latter, enlarged, showing the expanded tips or dises by which 


they cling. 


t 
dt 


- 
- 
~ 


SPECIAL FORMS. 


the figure.’ 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 any solid surface, such as a wall 
or tree-trunk, the tip expands into an adhesive disc, which forms 
a secure attachment. (Fig. 94.) Ina related plant, Vitis (Cissus) 
tricuspidata of Japan, these disks terminate the branches of very 
short tendrils: consequently 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. 
N 


111. The simple tendril of a Passion- 
flower, being in the axil of a leaf (that 
isin the position of a branch), is also of 
axial nature: it is a leafiess 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 
sicyoides. 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 
‘= =owhich 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 stem or branch, indurated, leafless, and 
attenuated to a pomt. 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 


1 This forms what is called a Sympodium or Sympodial stem, which is mor- 
phologically made up of a series of superposed branches. (See 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. 


56 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- 
cessory buds, placed far above 
the axil, that develops into 
the thorn (Fig. 96). Here the 
spine itself usually branches, 
and sometimes becomes ex- 
tremely compound. 

115. 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 
superiicial outgrowths from 
the bark, of? a different nature 
(385), and of small morpho- 
logical signification. 

114. Subterranean Stems are hardly less diverse than the 
aerial. They are classed as Ruizomes, Tusers, Corms, and 
Buss, the forms passing one into another by gradations. 

115. Rhizoma (/hizome, or in English Roorsrocxk) 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 coy- 
ered by the soil, and usually produces from its apex some kind 
of aerial stem, either leafy or as a flower-stalk (scape, 97), 
which rises into the air and sight. . Before morphology was 
understood, rootstocks were called creeping roots, scaly roots, &e. 
Some are slender, such as those of Mints (Fig. 99), of most 
Sedges (Fig. 98), and of Couch-Grass. Their cauline nature is 
evident from their structure and appearance; their nodes and 
internodes are well marked, the former bearing leaves reduced to 


FIG. 96. Branching thorn of the Honey-Locust (Gleditschia), an indurated branch 
developed from an accessory bud produced above the axil. a. Three buds under the 
base of the leaf-stalk, brought to view in a section of the stem and leaf-stalk below. 

FIG. 97. Thorn of the Cockspur Thorn, developed from the central of three axillary 
buds; one of the lateral buds is seen at its base. 


SPECIAL FORMS. 


ay 
t 


7 


seales , and the advancing apex rises at length into an ordinary 
stem, while the opposite and older end gradually dies away. A 
bud forms in the axil of each seale-like leaf, or 
in some of them; roots proceed from the nodes 
in preference ; the destruction of the ascending 
Stem only brings these buds into activity ; and 
the cutting or tearing of the rootstock into 
pieces by the hoe or plough merely hastens the 
establishment of as many new plants, each with 
roots, bud, and a small store of nourishment 
ready provided. It is this which makes Couch- 
Grass or Quick-Grass (Triticum repens) very 


a 


ees 


98 


troublesome to the agriculturist; and the Nut-Grass (Cyperus 
rotundus, var. Hydra) of the Southern Atlantic States is even 


more so, portions of its rootstock being tuberiferous, ¢. e. en- 
larged into a tuber which contains a supply of 
concentrated nourishment to feed the growth. 
116. Thickened rootstocks are common ; 
nourishing matter, elaborated in the leaves 
. above, being accumulated in them, just as 
it is in thickened roots, and for the same pur- 
pose. (53-55.) Such are the so-called roots of Sweet-Flag, of 
Ginger, of Iris or Flower-de-Luce (Fig. 216), of Bloodroot, of 
Solomon’s Seal (Fig. 100), &c. These grow after the manner 
of ordinary stems, advancing from year to year by the annual 
development of a bud at the apex, and emitting roots from the 
under side or the whole surface. Thus established, the older 


FIG. 98. Slender rhizoma of Carex arenaria, of Europe, which binds shifting sands 
of the sea-shore. 

FIG. 99. Rootstocks, or creeping subterranean branches, of the Peppermint. 

‘FIG. 99. A piece of the rootstock of the Peppermint, enlarged, with its node or joint, 
and two axillary buds ready to grow. 


58 MORPHOLOGY OF STEMS. 


portions die and decay as corresponding additions are made to 
the opposite growing extremity. -Each year’s growth is often 


marked conspicuously, sometimes by a strong contraction where 
the interruption took place, as in certain species of Iris (Fig. 216); 

or by the circular im- 

pressed scar (likened to 

the impression ofa seal) 
* jin Solomon’s Seal; this 
ye, being the place where 
J the annual aerial stem, 
bearing the vegetation, 
separated in autumn 
from the perennial rhi- 
zoma. The numerous 
slender lines encircling the rootstock are the scars left after the 
decay of the scale-like leaves or bud-seales, such as are seen at 
4 the young and growing end of the rootstock. 
The rootstock of Diphylleia, of the Alleghany 
Mountains (Fig. 101), is similar; but the 
yearly growths are so exceedingly short that 
they become vertical, the bud of each year, 
is close to the stalk of the year preceding, 
and the scarsanarking preyious growths are 
in contact.? villi makes a short and 
mostly vertical rootstock, which, when it 
remains simple and dies away promptly 
below (as in Fig. 102), comes nearly within 
the definition of a corm. But in several 


1 The rootstock in Polygonatum and Diphylleia is a sympodium (110, note), 
the terminal bud developing yearly the growth above ground and perishing 


FIG, 100. Rootstock of Polygonafum or Solomon’s Seal, with the terminal bud, the 
base of the stalk of the season, and three scars from which the latter has separated in 
as many former years. 

FIG. 101. Rhizoma of Diphylleia eymosa, showing six years’ growth, and a bud for 
the seventh: a, the bud: 6, base of the stalk of the current year: c, scar left by the © 
decay of the annual stalk of the year before; and beyond are the scars of previous years. — 

FIG. 102. Shoot and young rootstock of Trillium erectum, with only terminal bud. 


3 


9 


ube 


SPECIAL FORMS. 


species, and in older individuals, it is longer, often oblique, and 
branching, and bears the scars from which the annual aerial 
growths have separated.*/ Nymphzea odorata, 
the sweet-scented white Water Lily, grows by 
very long, stout, and simple rootaiagl es: ) in 
NW; tuberosa the sides of the. rootstag 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 involy- 
ing several nodes —is thickened by the depo- 
sition of nourishing matter. A potato and a 
Jerusalem artichoke are typical 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- B 
iferous branches 
may be __ in- 
creased. The number of nodes and internodes involvea 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 spec ies, 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. ) ~ 


a 
. 


GU MORPHOLOGY OF STEMS. 


be called a Monomerous tuber, namely in Nelumbium luteum 
(Fig. 1087), where it consists of a single thickened internode of 


107 106 108 


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 
1037 are strung as it were upon’ 
a long filiform axis), the tubers not unfrequently being mo- 
nomerous, although the larger ones are not so. 
117°. Tubereles, as they may be termed, are of a a 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. 106. Forming potatoes in various stages. 107. One of the younger ones en- 
larged. 108. Section of a small portion passing throngh an eye, or bud, more enlarged. 

FIG. 108¢, 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,! 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.” 

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 alla. I 
the vegetation and the blossoms of the season. if KS i z 
A good type of corm is that of Cyclamen LAN 
(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 corm of Indian 
Turnip (Ariszema triphyllum, 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. 111, 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 bu/bs. Inecommon parlance, they 
will doubtless continue to be called bulbs, and even in popular 


1 Not, however, such as those of Aplectrum, Tipularia, etc., which are 
genuine corms or tubers. = 

* Irmisch, Beitr. Biol. & Morphol. Orchid. 1853, fide Duchartre, Elém. 
Bot. 278. 


FIG. 109. Depressed corm of Cyclamen. 
FIG. 110. Corm of Indian Turnip, Arisema triphyllum. 


62 MORPHOLOGY OF STEMS. 


botanical descriptions. In fact, while they differ from naked 
corms in having some investment, they differ from true bulbs 
only in the greater size of the solid axis and the fewness of 
the investing scales; 

the stem or solid body 
making the greater part 
of the corm, but a very 
small part of a proper 
bulb. There are, more- 
Ny, over, all gradations be- 
11 112 tween the two. 

120. 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 leayes. 
Compared with buds (73), it is a very fleshy bud, usually large 
and subterranean, the axis of which never elongates. It is a 


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, théfew thin enveloping scales removed, showing their 
scars, which mark the nodes, the shrivelled vestige of the last year’s corm at the base, 
and buds developing into new ones on various parts of its surface. 112. Vertical section 
of a similar corm, with a terminal and one lateral bud. 

FIG. 113. Section of a tunicated bulb of the Onion. 

FIG. 114. Vertical section of the bulb of the Tulip, showing its stem or terminal 
bud (c) and two axillary buds (8, 5). 

FIG. 115. Bulb of a Garli:, with a crop of young bulbs. 

FIG. 116. Vertical section of the corm of a Crocus: a. new buds. 

FIG. 117. Vertical section of the corm of Colchicum (6), with the withered corm of 
the preceding (a), and the forming one (c) for the ensuing year. 


SPECIAL FORMS. 63 


121. A Tunicated or Coated Bulb (Fig. 113-115) is one in 
which the scales are broad and completely enwrapping, forming 
concentric coatings. These are thickish when fresh, but thin 
when exhausted and dry, as in the Onion, Garlic, and Tulip. 

122. A Sealy Bulb 
has the bulb-scales 
comparatively —nar- 
row, thick, and small, 
imbricated, but not 
severally enwrapping 
each other. That of 
the Lily is the mest | 
familiar and char- Wd) 
acteristic example. 
Mie) 118; 119:) 

123. Bulblets are small aerial bulbs, or buds with fleshy scales, 
which arise in the axils of the leaves of several plants, such as 
the common Lilium bulbiferum and 
L. tigrinum, the Tiger Lilies of the 
gardens (Fig. 120). Here they ap- 
pear during the summer as axillary 
buds: they are at length detached, 
and falling to the ground strike root, 
and grow as independent plants. In 
the common Onion, and in many other 
species of- Allium, similar bulblets 
take the place of flower-buds in the 
umbel. Bulblets plainly show the identity of bulbs with buds. 

124. All these extraordinary, no less than the ordinary, forms 
of the stem, grow and branch, or multiply, by the development 
of terminal and axillary buds. This is perfectly evident in the 
rhizoma and tuber, and is equally the case in the corm and bulb. 
The stem of the bulb is usually reduced to a mere plate (Fig. 
114 4a), which produces roots from its lower surface, and leaves 
or scales from the upper. Besides the terminal bud (c), which 
usually forms the flower-stem, lateral buds (4, 6) are produced 
in the axils of the leaves or scales. One or more of these may 
develop as flowering stems the next season, and thus the same 
bulb survive and blossom from year to year; or these axillary 
buds may themselves become bulbs, feeding on the parent bulb, 
which in this way is often consumed by its own offspring, as in 


=) 


FIG. 118. Sealy bulb of Canada Lily, Lilium Canadense, after flowering. 119. Ver- 
tical section of same, showing two new young bulbs within. 
FIG. 120. Bulblets in the axil of the cauline leaves of Tiger Lily. 


64+ MORPHOLOGY OF STEMS. 


the Garlic (Fig. 115); or, finally separating from the living 
parent, just as the bulblets of the Tiger Lily fall from the stem, 
they may form so many independent individuals. So the corm 
of the Crocus (Fig. 111, 112) produces one or more new ones, 
which feed upon and exhaust it, and take its place; and the 
next season the shrivelled remains of the old corm may be found 
underneath the new. The corm of Colchicum (Fig. 117) pro- 
duces a new bud on one side at the base, and is consumed by it 
in the course of the season; the new one, after flowering by its 
terminal bud, is in turn consumed by its own offspring; and 
so on. The figure represents at one view, a, the dead and 
shrivelled corm of the year preceding; 6, that of the present 
season (in a vertical section); and, ¢, the nascent bud for the 
growth of the ensuing year. 

125. Condensed Stems, homologous with corms, tubers, &e., 
and similar in mode of growth, but above ground, and multiply- 
ing in the same ways, are not uncommon. The Cactus family is 
mainly composed of such forms, of flat- or round-jointed Prickly 
Pears (Opuntia), fluted or angled columns (Cereus), and glob- 
ular Melon-Cactus, Mamillaria, and Echinocactus. The latter 
types, which completely imitate corms, are the most consolidated 
forms of vegetation. While ordinary plants are constructed on 
the plan of great expansion of surface, these present the least 
possible amount of surface in proportion to their bulk, their 
permanent spherical figure being that which exposes the smallest 
portion of their substance to the air. Such plants are evidently 
adapted to very dry regions ; and in such only are they naturally 
found. Similarly, bulbous and corm-bearing plants, and the 
like, are a form of vegetation which in the growing season may 
in the foliage expand a large surface to the air and light, while 
during the period of rest the living vegetable is reduced to a 
globular or other form of the least surface ; and this is protected 
by its outer coats of dead and dry scales, as well as by its subter- 
ranean situation ;— thus exhibiting another and very similar 
adaptation to a season of drought. And such plants mainly 
belong to countries (such as Southern Africa, and the interior 
of Oregon and California) which have a long hot season, during 
which little or no rain falls, when, their stalks and foliage above 
and their roots beneath being early cut off by drought, the plants 
rest securely in the corm-like forms to which they are reduced, 
and retain their moisture with great tenacity until the rainy season 
returns. Then they shoot forth leaves and flowers with wonderful 
rapidity, and what was perhaps a desert of arid sand becomes 
green with foiiage and gay with blossoms, almost in a day. 


SPECLAL FORMS. 65 


126. Stems serving the purpose of foliage, Phyllocladia. Most 
of these condensed and permanent stems are illustrations of 
this, their green rind doing duty for leaves, which 
are either absent, or transient, or reduced to 
spines or other organs not effective as foliage. 
In the flat and broad-jointed species of Opuntia, 
and still more in Phyllocactus and Epiphyllum, the 
forms assumed give a considerable surface of green 
rind, which well answers the purpose of leaves. 
Flattened stems or branches of the same sort and 
economy not seldom occur in other than fleshy or 
succulent plants (such as the Cactuses) ; some- 
times accompanied by a certain number of real 
foliage-leaves, but these more or less transient, 
as in Bossiza and Carmichelia among Legu- 
minous shrubs, and Muhlenbeckia platyclada, now 
in common cultivation (Fig. 121); sometimes 
with all the leaves reduced to small and function- 
less scales, as in the Xylophylla (7. e. wooden- 
leaved) section of Phyllanthus, and in Phyllo- 
cladus (New Zealand and Tasmanian trees of 
the Yew family). In all these, the cauline nature 
is manifest by the continuous or proliferous 
growth, by the marked nodes and internodes, 
and often by the bearing of flowers. 

127. Cladophylla (literally, branch-leaves) are more ambigu- 
ous in character. The most familiar examples are found in the 
peculiar foliage of Ruscus, Myrsiphyllum, Asparagus, and in 
some other genera of the same family. In these the primary or 
proper leaves of the shoots are little scales, one to each node, and 
quite functionless. From the axil of each is immediately pro- 
duced a body answering in all respects to the blade of a leaf, 
both in appearance and in office. They also accord with leaves 
in being expanded horizontally, although they take a twist which 
brings them more or less into a vertical position, in the manner 
of phyllodia (that is, of leaf*stalks assuining the form and office 
of leaf-blades, 217); wherefore they may be regarded as the 
first and only leaf of an axillary branch with the internode 
under the leaf wholly undeveloped and no further growth ever 
taking place. But, on the other hand, their anatomical structure 
is said to be that of stems rather than of leaves. Moreover, 


FIG. 121. Foliiform branch of Mublenbeckia platyclada. growing from the apex, 
bearing a smal! and transient leaf at some nodes, also a fluwer or two. 


66 MORPHOLOGY OF STEMS. 


the cladophyll of Ruscus (called Butcher’s Broom in England, 
Fig. 123) not only becomes firm, hard, and spiny-tipped, but 
it exhibits the character of a 
branch by bearing flowers on the 
middle of one face, in the axil of 
a little bract. Under this view 
such a cladophyll would seem to 
be a flattened branch of two in- 
ternodes, or else of one internode 
with a flower-stalk adnate to it. 
In Myrsiphyllum (a South A fri- 
can climber, commonly cultivated 
under the erroneous name of 
Smilax, Fig. 122), the cladophyll 
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, Bischoff has 
given the name PHYLLocLapiA, sing. PHytLoctapium. To those 
definitely restricted to one internode, and which so closely 
counterfeit leaves, Kunth gave the name of CLapopra, sing. 
Ciapopium. 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 
was 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 described. But for those of 
the preceding paragraph, which are so peculiarly 
leaf-like, Kunth’s name of Cladodium (t. e. a 
branch-like body) is false in meaning, and may 
be replaced by that of CLApopiyLLuM (7. e. leaf- 
branch), or in shorter English CLapopnyLt. 

129. Frondose Stems. Finally, in some few 
124 pheenogamous plants, the whole vegetation is re- 
duced toa simple leaf-like expansion, as in Duckweed (Lemna), 


122 123 


FIG. 122. Myrsiphyllum, with cladophylls serving for foliage; the true leaves eon- 
sisting of minute and very inconspicuous scales subtending the former. 

FIG, 123. Asingle cladophyll of 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. Lemna minor, a common Duckweed, whole plant in flower, magnified. 


INTERNAL STRUCTURE. 67 


Fig. 124. Here is no differentiation whatever into stem and 
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 body, which answers both for 
stem and foliage, is termed a Fronp, 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 phanogamous 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. 

13i. The stems of phzenogamous 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. Bothare equally composed of cells, or origi- 


cs 


a A 
Hat oa ae 


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 


Jee Se EE EE ee eee eee ee eee eee 


PIG. 124%. A magnified slice of a portion of the flower-stalk of Richardia ZEthiopica 
(the so-called Calla Lily), transverse with some longitudinal view: mainly parenchyma, 
the vells built up so as to leave comparatively large vacancies (intercellular spaces or 
alr-passages); near the centre a cross-section of a fibro-vascular bundle, and next the 
margin or rind some finer ones. 


638 MORPHOLOGY 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 


000 uo9 


0) O00 
0 


a> 


092 920 QO 


VON 0 


ung 


WOOUOCD.O0 0 6 


hw) 


Uo 


CCK 


= 
EC 
Zg 


(Cauca 
(( 


Cac 


LIC 


c 
125 


short and commonly thin-walled cells, form cellular tissue. Its 
ordinary form (of roundish, cubical, or polyhedral and thin-walled 
cells) 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-vyasecular elements. a. Bast-cells (long wood-cells) of fibrous bark 
of Linden or Bass-wood. b. Some wood-cells and (below) a duct, and ¢ a detached 
wood-cell of the wood of same tree, equally magnified with a. d. A detached wood- 
cell from a shaving of White Pine, showing the peculiar disk-like markings. e. Portion 


of same shaving. jf. Portion of a dotted duct from the Vine, evidently made up of a 
series of short cells. g. Part of a smaller dotted duct, showing no appearance of such 
eomposition. hk, i. Spiral ducts or vessels, of the ordinary kind. j. Spiral duct of 
Banana. k. Duct from Celery, the thread within spiral or annular below, reticulated 
above, and higher passing into the state of dotted duct. /. Duct from Impatiens, with 
the open spiral passing into rings at the middle. All magnified 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. 

132. In the forming state, the whole stem is parenchyma ; 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 phenogamous plants, 
which haye been termed the Endogenous and the Exogenous,’ 
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.” 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 year’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 jj) 
endogenous stem has the wood in distinct : i i 
threads or fibro-yascular bundles, travers- § | | | 
ing the cellular system or parenchyma with 9) | | 
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 DeCandolle, following the ideas of Desfontaines, 
and which have played an important part in structural and systematic 
botany ever since DeCandolle adopted these names as those of the two 
primary divisions of phenogamous plants, Lxogene and Endogene. 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, Monocoiyledones 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. 

2 Yet with some more or less valid exceptions, as when the annual stem 
of Podophyllum and the rhizoma of Nymphza, among dicotyledonous plants, 
imitate the endogenous structure; or where the pith of an evidently exogenous 
stem, as in the Piperacew, has scattered woody bundles in an endogenous 
fashion ; or where monocotyledonous plants have all their woody bundles in 
a definite circle, as in Luzula, Croomia, &c. 

FIG. 126. Se tion 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 Bamboo, 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 yari- 
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 only 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 ENpoGENous 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 Palms —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 Chamerops 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 unbranched 
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 
erow 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 species, regularly 
in the Doum Palm of Upper Egypt, and in the Pandanus, or 
Screw-Pine (Fig. 69), which belongs toa 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 
the Bamboo, Maize, 
&ce.), they gradually 
taper upward in the 
manner of most ex- 
ogenous stems. 

for. “This~ kind 
of stem comprises 
several subordinate 
types as to internal 


('X 


ry aN SP = 


: a 
structure, which to a 
be well understood ba 
must be studied his- a 
tologically, under the Ea 


ae 
Hf 


pase 


microscope.’ Toone 
of these, by no means 
the simplest, belongs 
the ordinary palm- 
stem, the anatomy 
of which was made 


classical by Mobhl, 3 
and has been 5 3 

wi ¢ 
supplemented \ N\ Ay , : 
by Negeli. 9 ey WZ i 
In this a large = \lf \ Ki 


part of the =Zsthy 
bundles, or all 7). 

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 l’Anatomie comparée et le Développement des 
Tissus de la Tige dans les Monocotylédones, 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. 


72 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 
4 the wood and rind, or no proper rind at 
Hall. In others, such as the Palmetto 
4 (Fig. 126), there is a marked rind 
or false bark, which receives independent 
fibro-vascular bundles from the leaf-stalks, 
and is traversed by them in parallel lines. 
oq In Grass-stems, and others with long inter- 
ib i nodes and closed nodes, the fibro-vascular 
C/iis bundles all run approximately straight 
: and parallel through the internodes, but are 
jij intricate and anastomosed in the nodes. 
5d The whole centre of the internodes, 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 
Nyctaginaceze and some Araliacee. Endogenous stems of 
simpler structure, as in herbaceous Liliaceze, Commelynacez, 
&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 Uvularia 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. 
138. An annual endogenous stem increases in diameter by 
general growth until it attains its limit. Ligneous and enduring 
stems increase similarly up to a certain period. Then the rind 


WN ll 


{ ) 


FIG. 127. 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 Draczena (Dragon-trees), in the arborescent Yuccas, and the 
like, the zone intermediate between the cortical and interior re- 
gion, which is for a time 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 haye 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 by 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, f-/), 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, b, 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 has formed a com- 
plete circle, interrupted only by the medullary rays, which radiate from the pith to 
the bark. 


74 MORPHOLOGY OF STEMS. 


surround a purely cellular central part (the pith), while sur- 
rounded by a cellular external rind, the bark, or outer bark. 
The diagrams (Fig. 128-150) rudely show some stages in the 
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 ifto the forming stem. 

142. First Year’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 cel- 
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 Mrp- 
ULLARY Rays. They 
represent the cellular 
system of the wood it- 
self, or untransformed 
parenchyma. Being 
pressed by the woody 


of 


PUN CCU CHP CHECK Cer eLE 


tt 


EF, 
ssa 


RECRETTC TCC 


LK 
Pe wedges, their cells are 
Rene cit va e  «¢ aif  @ pe laterally fattemedsssam 
38 some stems, the med- 


ullary rays, or many 
of them, are comparatively broad and conspicuous; in others, 
thin 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- 


FIG. 131. Longitudinal and transverse section of a stem of the Soft Maple (Acer 
dasycarpum), at the close of the first year’s growth; of the natural size. 

FIG. 132. Portion of the same, magnified, showing the cellular pith, surrounded by 
the wood, and that by the bark. 

FIG. 133. More magnified slice of the same, reaching from the bark to the pith: 
a. part of the pith; 0. vessels of the medullary sheath; ec. the wood; d, d. dotted ducts in 
the wood; ¢, e. annular ducts; 7. the liber, or inner fibrous bark; g. the cellular envelope, 
or green bark; h. the corky envelope; 7. the skin or epidermis; &. one of the medullary 
rays, seen on the transverse section. 


INTERNAL STRUCTURE. i) 
tomical structure of a woody exogenous stem of a year old is 
displayed in the Fig. 131-135. Viewing the parts particularly, 
and in order from centre to circumference, there is, — 

Ist. The Pith or Medulla, consisting entirely of soft and rather 
large thin-walled cells,* gorged with sap or other nourishing 
matter during the growing state, becoming light, dry, and empty 
when effete. 

2nd. The Layer of Wood, traversed by the medullary rays. In 
Pines and other Conifers, the wood is of uniform structure, being 
wholly composed of a woody tissue with peculiar markings (Fig. 
125, d,e) : in other wood, ducts of one or more sorts occur ; the 
most conspicuous being what are termed dotted ducts. These 
are so large as to be evident to the naked eye in many ordi- 
nary kinds of wood, especially where they are accumulated in 
the inner portion of the layer, as in the Chestnut and Oak. In 
the Maple, Plane, &c., they are rather equably scattered through 
the annual layer, and are too small to be seen by the naked eye. 
Next the pith, 7. e. in the very earliest formed part of the wood, 
some spiral ducts are uniformly found, and this is the only part 
of the exogenous stem in which these ordinarily occur. They 
may be detected by breaking a woody twig in two, after dividing 
the bark and most of the wood by a circular incision, and then 
pulling the ends gently asunder, when their spirally coiled fibres 
are readily drawn out as gossamer threads. As these spiral 
ducts form a circle immediately surrounding the pith, they have 
collectively been termed the medullary sheath, but they hardly 
deserve a special name. ‘The vertical section in Fig. 153 divides 
one of the woody wedges, and shows no medullary ray ; but there 
is one at the posterior edge of the transverse section. But, in the 
much more diagramatic Fig. 154, the section is made so as to show 
the surface of one of these plates, or medullary rays, passing hori- 
zontally across it, connecting the pith (p) with the bark (4). 
These medullary rays form the silver-grain (as it is termed), which 
is sO conspicuous in the Maple, Oak, &c., and which gives the 
glimmering lustre to many kinds of wood when cut in this direc- 
tion. _A section made as a tangent to the circumference, and 
therefore perpendicular to the medullary rays, brings their ends 
to view, as in Fig. 155, much as they appear on the surface of a 
piece of wood from which the bark is stripped. They are here 
seen to be composed of parenchyma, and to represent the horizon- 


1 In rare instances, a few fibro-vascular threads are found dispersed 
through the pith, presenting a somewhat remarkable anomaly. This 
occurs in Aralia racemosa, and more strikingly in Mirabilis and other 
Nyctaginacee, and in Piperacexw. (133, foot-note.) 


76 MORPHOLOGY OF STEMS. 


tal system of the wood, or the woof, into which the vertical woody 
fibre, &c., or warp, isinterwoyen. The inspection of a piece of oak 
or maple wood at once shows the pertinency of this illustration. 


i 


hel 


ti 
i 


135 


3rd. The Bark or rind. This at first consisted of simple 
parenchyma, like that of the pith, except for the green color 
developed in it, the same as that which gives verdancy to foliage. 
This green matter is formed in the cells of all such parts when 
exposed to light, consists of green grains of somewhat complex 
chemical composition, has important functions to perform in 
assimilation (¢.e. in the conversion of the plant’s crude food into 
vegetable matter), and is named CHLOROPHYLL, 7. e. leaf-green. 
The completed bark, when all its parts are apparent, as espe- 
cially in-most trees and shrubs, is composed of three strata, of 
which the green bark, the most conspicuous in the young shoot, 
is the middle layer, therefore named the MesopnHtaum. This is 
soon covered, and the green color obscured, by a superficial 
stratum of cells, generally of some shade of ash-color or brown, 
occasionally of brighter tints, which gives to the twigs of trees 
and shrubs the hue characteristic of each species, the CorKxy 
EnvVeELore or layer, or EpipHta@um. The latter name denotes its 
external position; the former, that it is the layer which, when 
much developed, forms the cork of Cork-Oak and those corky 
expansions which are so conspicuous on the twigs of the Sweet 
Gum (Liquidambar), and on some of our Elms (Ulmus alata 
and racemosa). It also forms the paper-like exfoliating layers 
of Birch-bark. It is composed of laterally flattened parenchy- 
matous cells, much like those of the Eprpermuis (Fig. 133, 2), 
which directly overlies it, and forms the skin or surface of the 


FIG. 134. Vertical section through the wood of a branch of the Maple, a year old, 
so as to show one of the medullary rays, passing transversely from the pith (p) to the 
bark (6): magnified. But a section can seldom be made so as to show one unbroken 
plate stretching across the wood, as in this instance. 

FIG. 135. A vertical section across the eucs of the medullary rays: magnified. 


T7 


INTERNAL STRUCTURE. 


stem, and of the whole plant. Lastly, the inner bark, accord- 
ingly named EnporHievm, takes the special name of Liser, 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 Basswood, contains two peculiar kinds of cells in 
BS C Ww 


cea) 


QLO<e 
tienes 


aan 
oa 
50 
vs) 


1354 mr 
addition to common parenchyma, both of the fibrous or vascular 
class: viz., 1. Criprirorm or StevE-cELLs, a sort of ducts the 
walls of which have open slits, through which they communicate 
with each other; 2. Bast or Bast-cexis, the fibre-like cells 
which give to the kinds of inner bark that largely contain them 


FIG. 135¢ Portion of a transverse section (above), and a corresponding vertical see- 
tion (below), magnified. reaching from the pith (p) 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 unrollable ducts, one seen length- 
wise with uncoiling extremity in the lower figure: w, w. woody tissue: dd. one of the 
dotted ducts interspersed in the wood: c/. carnbium-layer or zone of new growth of 
wood and inner bark: /-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. 


(9 6) 


7 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 greatly exceeds the calibre. This is the 
material which gives to 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 year’s growth, and immedi- 
ately before that of the second year begins, see Fig. 135%.) 

143. Annual Increase in Diameter. An herbaceous stem does 
not essentially differ from a woody one of the same age, except 
that the wood forms a less compact or thinner zone; ‘and the 
whole perishes, at least down to the ground, at the close of the 
season. Buta 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, by 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 Camprum-Layer. It is charged with 
organizable matter, which is particularly abundant and mucila- 
ginous in spring when growth recommences. This mucilaginous 
matter was named Cambium 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 heowern 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 off 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 w ood 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.? 

144. Demareation 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 
eye, 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 woody 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 cells, thus marking 
an 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. 

145. Each layer of wood, once formed, remains essentially 
unchanged 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. 

146. Sap-wood (Atsurnum). In the plantlet and in the 
developing bud, 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 bears 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 
woody trunks the sap continues to rise year after year 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 
layers 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 the fresh rootlets that imbibe with the 
foliage that elaborates the sap, is each year removed farther from 
them. The latter, therefore, after a few 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 abrupt and conspicuous —into sap-wood and hear-wood. 
The former is the popular name given to the outer and newer 
layers of softer, more open, and bibulous wood. The early physi- 
ologists named it alburnum from its white or pale color. Being 
more or less sappy, or containing soluble organic matter, and 
readily imbibing moisture, this part of the wood is liabie to decay, 
and it is therefore discarded from timber used for construction. 

147. Heart-wood (or DurAmey, so called from its greater hard- 
ness or durability) is the older and mature portion of the wood. 


INTERNAL STRUCTURE. 81 


In all trees which have the distinction between the sap-wood and 
heart-wood well marked, the latter acquires a deeper color, and 
that peculiar to the species, such as the dark brown of the Black 
Walnut, the blacker color of the Ebony, the purplish-red of Red 
Cedar, and the bright yellow of the Barberry. These colors are 
owing to special vegetable products, or sometimes to alterations 
resulting from age. In the Red Cedar, the deep color belongs 
chiefly to the medullary rays. In many of the softer woods, there 
is little change in color of the heart-wood, except frem incipient 
decay, as in the White Pine, Poplar, Tulip-tree, &c. The 
heart-wood is no longer in any sense a living part: it may perish, 
as it frequently does, without affecting the life or health of the 
tree. 

148. The Growth and Duration of the Bark, also the differences 
in structure, are much more various than of the wood. Moreover, 
the bark is necessarily subject to grave alterations with advanc- 
ing age, on account of its external position; to distention from 
the constantly increasing diameter of the stem within, and to 
abrasion and decay from the influence of the elements without. 
It is never entire, therefore, on the trunks of large trees; but 
the dead exterior parts, no longer able to enlarge with the en- 
larging wood, are gradually fissured and torn, and crack off in 
strips or pieces, or disappear by slow decay. So that the bark 
of old trunks bears only a small proportion in thickness to the 
wood, even when it makes an equal amount of annual growth. 

149. The three parts of the bark (142), for the most part 
readily distinguishable in the bark of young shoots, grow inde- 
pendently, each by the addition of new cells to its inner face, so 
long as it grows at all. The green layer commonly does not 
increase after the first year ; the opaque corky layer soon excludes 
it from the light; and it gradually perishes, never to be renewed. 
The corky layer usually increases for a few years only, by the 
formation of new tabular cells : occasionally it takes a remarkable 
development, forming the substance called Cork, as in the Cork 
Vak, and the thin and parchment-like layers of the White and 
Paper Birches. 

150. The liber, or inner bark, continues its growth through- 
out the life of the exogenous tree, by an annual addition from 
the cambium-layer applied to its inner surface. Sometimes this 
growth is plainly distinguishable into layers, corresponding with 
or more numerous than the annual layers of the wood: often, 
there is scarcely any trace of such layers to be discerned. In 
composition and appearance, the liber varies greatly in different 
plants, especially in trees and shrubs. That of Basswood or 


82 MORPHOLOGY OF STEMS. 


Linden, and of other plants with a similar fibrous bark, may be 
taken as best representing the liber. Here it consists of alter- 
nate strata of fibrous bast, and of the peculiar liber-cells called 
sieve-cells, in which nourishing matter is especially contained 
and elaborated. While the latter, or their equivalents, occur 
and play an important part in all inner bark, the bast-cells are 
altogether wanting in the bark of some plants, and are not pro- 
duced after the first year in many others. The latter is the case 
in Negundo, where abundant bast-cells, like those of Basswood, 
compose the exterior portion of the first year’s liber, but none 
whatever are formed in the subsequent layers. In Beeches and 
Birches, also, a few bast-cells are produced the first year, but 
none afterwards. In Maples, a few are formed in succeeding 
years. In the Pear, bast-cells are annually formed, but in very 
small quantity, compared with the parenchymatous part of the 
liber. In Pines, at least in White Pines, the bark is nearly as 
homogeneous as the wood, the whole liber, except what answers 
to the medullary rays, consisting of one kind of cells, resembling 
those of bast or of wood in form, but agreeing with the proper 
liber-cells in their structure and markings. 

151. The bark on old stems is constantly decaying or falling 
away from the surface, without any injury to the tree; just as 
the heart-wood within may equally decay without harm, except 
by mechanically impairing the strength of the trunk. ‘There are 
great differences as to the time and manner in which the older 
bark of different shrubs and trees is thrown off. Some have 
their trunks invested with the liber of many years’ growth, 
although only the innermost layers are alive; in others, it scales 
off much earlier. On the stems of the common Honeysuckle, of 
the Nine-Bark (Spirzea opulifolia), and of Grape-vines (except 
Vitis vulpina), the liber lives only one season, and is detached 
the following year, hanging loose in papery layers in the former 
species, and in fibrous shreds in the latter. 

152. 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 convey 
to the cambium-layer or zone of growth. The proper juices and 
peculiar products of plants are accordingly found in the foliage 
and the bark, especially in the latter. Im the bark, therefore 
(either of the stem or of the root), medicinal and other principles 
are usually to be sought, rather thanin 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 deveiop the foliage. 2d, The fresh roots and rootlets 
annually developed at the opposite extremity. 35d. The newest 
strata of wood and bark, and especially the interposed cambium- 
layer, 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 theSap 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 laver 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 years, and some are possibly 
older.1 Equally long may survive such endogenous trees as the 

Dragon tree (Draczena), 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 way 
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 “ Physiologie Végétale ;” 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. 


8+ MORPHOLOGY OF STEMS. 


which the aged are either increasingly incident or less able to 
resist. A tree like the Banyan (59, Fig. 71), which by aerial 
roots continues to form new trunks for the support and sustenance 
of the spreading branches, 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.’ However this may be, it is evident that 
a 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 ;? that rather 

156. The Plant is a Composite Being, or Community, lasting, in 
the case of a treg, through an indefinite and often immense num- 
ber of generations. These are successively prodaced, 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. No 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 mternodes 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 offspring 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 many 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. 


MORPHOLOGY 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. Or Leaves. 
§ 1. THerr NATURE AND OFFICE. 


157. Leaf (Lat. Yolium, 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 way 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, BLApE or 
Lamia; 2, Foor-statk or Lear-sraLk, technically PErio.e ; 
3, A pair of Stieutes. (Fig. 142.) 

158. The blade, being the most important part 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 the true nature of organs. 

160. Morphologically, and in the most comprehensi¥e 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-bulb 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 jfoliage-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 
phylline 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... Whatever is produced in the axil of a leaf is 
cauline, and when developed is a branch. 

162. 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 leafless, 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 There are cases in which this rule is of difficult application, or is seem- 
ingly violated, sometimes by the suppression of the subtending leaf, as in the 
inflorescence of Crucifere, rarely in other ways, to be explained in the 
proper places. 


none 


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 they all perish in autumn nearly 
simultaneously. 

163. Leaves soon complete their growth, and have no power 
of further increase. Being organs for transpiration, a very large 
part of the water imbibed by 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 Palms, 
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. (215-217.) 


§ 2. THerr Structure AND Forms AS ORGANS OF ASSIMILATION 
orn VEGETATION, @. 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- 


85 MORPHOLOGY OF LEAVES. 


work, the separate parts or ramifications of which form what are 
variously called the 77s, — a sufticiently proper term, — nerves or 
veins. ‘The latter names may suggest false analogies ; but they 
are of the commonest use in descriptive botany. That of veins, 
and of its diminutive, vein/ets, for the smaller ramifications, is 
not amiss; for the fibrous framework not only gives firmness 
and support to the softer cellular apparatus, 7. e. forms ribs, 
but serves in the leaf, as it does in the stem, for the more rapid 
conveyance and distribution of the sap. The subdivisions con- 
tinue beyond the limits of unassisted vision, until the fibro-vas- 
cular bundles are reduced to attenuated fibres ramified through 
the parenchyma. In leafstalks, the woody bundles are parallel, 
not ramified, and arranged in various ways; in Exogens usually 
so as to form in cross-section an arc or an incomplete or com- 
plete ring. In leaves serving as foliage or organs of assimilation, 
the blade is the important part, and this only is here regarded. 

168. The characteristic contents of these cells of parenchyma 
are grains of chlorophyll (142°), literally leaf-green, to which 
the green color of foliage is wholly owing, and which may be 
regarded as the most important of all vegetable products ; 
because it is in them (or in this green matter, whatever its form, 
189) that all ordinary assimilation takes place. As it acts only 
under the influence of light, the expanded leaf-blade may be 
viewed as an arrangement for exposing the largest practicable 
amount of this green matter — the essential element of vegeta- 
tion — to the light and air. 

169. The Parenchyma-cells, constituting the green pulp, are 
as eal Gas can | | themselves arranged in accordance 

Eee yseepees With this adaptation. The upper stra- 
tum is mostly of oblong cells, compactly 
arranged in one or more layers, their 
longer diameter perpendicular to the 
surface. The stratum next the lower 
surface of the leaf consists of loosely 
arranged cells, with longer diameter 
usually parallel to the plane of the leaf, 
often irregular in form, and so disposed 
as to leave intervening sinuous air-spaces freely permeating all 


FIG. 136. A magnified section through the thickness of a leaf of Iicium Flori- 
danum, showing the irregular spaces or passages between the cells, which are small in 
the upper layer of the green pulp, the cells of which (placed vertically) are well com- 
pacted, so as to leave only minute vacuities at their rounded ends; but the spaces are 
large and copious in the rest of the leaf, where the cells are very loosely arranged: 
also the epidermis or skin of the upper (a) and of the lower surface of the leaf (0d), 
composed of perfectly combined and thick-walled empty cells, 


THEIR. STRUCTURE AND FORMS. 89 


that part of the leaf. (Fig. 136, 137.) Hence in good part the 
deeper green hue of the upper, and the paler of the lower face 
of leaves. 

170. Epidermis. The whole surface of leaves, as of young 
stems, is invested with a translucent membrane, composed of 
one or sometimes two or three layers of empty and rather- 
thick-walled cells. This is the skin or epidermis, which is so 
readily separable from the succulent tissue of such leaves as 
those of Stonecrop and other species of Sedum. It is of a single 
layer in the Illicium (Fig. 136) and Lily (Fig. 137); of as 
many as three in the firm leaf of the Oleander; is generally 


187 


hard and thick in such coriaceous leaves as those of Pittosporum 
and Laurustinus, which thereby the better endure the dry air of 
rooms in winter. 

171. Stomata or Breathing-pores.t| The epidermis forms a 
continuous protective investment of the leaf except where certain 
organized openings occur, the stomata. They are formed by a 
transformation of some of the cells of the epidermis ; and consist 
usually of a pair of cells (called guardian-cells) , with an opening 
between them, which communicates with an air-chamber within, 
and thence with the irregular intercellular spaces which permeate 
the interior of the leaf. Through the stomata, when open, free 
interchange may take place between the external air and that 


1 The technical name has been anglicized stomates, singular stomate, which 
has no advantage over the proper Greek, sing. sfoma, pl. sfomata. 
FIG. 137. A magnified section through the thickness of a minute piece of the leaf 


of the White Lily of the gardens, showing also a portion of the uuder side with some 
breathing-pores, stomata. 


90 MORPHOLOGY OF LEAVES. 


within the leaf, and thus transpiration be much facilitated. 
When closed, this interchange will be interrupted or impeded. 
The mechanism of stomata is somewhat 
recondite, and will be illustrated in the 
anatomical and physiological volume of 
this series. 

172. It is only when leaves assume 
a vertical or edgewise position that the 
stomata are in equal numbers on both 
faces of a leaf. Ordinarily. they occupy 
or most abound on the lower face, which 
is turned away from the sun; but in certain coniferous trees the 
reverse of this is true. In the Water Lilies (Nymphzea, Nuphar), 
and other leaves which float 
upon the water, the stomata all 
belong to the upper surface. 
Leaves which live under water, 
where there can be no evapora- 

a a va tion, are destitute, not only of 
stomata, but usually of a distinct epidermis also. The number 
of the stomata varies from 800 to about 170,000 on the square 
inch of surface in different leaves. In the Apple, there are said 
to be about 24,000 to the square inch (which is under the average 
number, as given in a table of 36 species by Lindley) ; so that 
each leaf of that tree would present about 100,000 of these 
orifices. The leaf of Dragon Arum is said to have 8,000 
stomata to a square inch of the upper surface, and twice that 
number in the same space of the lower. That of the Coltsfoot 
has 12,000 stomata to a square inch of the lower epidermis, 
and only 1.200 in the upper. That of the White Lily has 
from 20,000 to 60,000 to the square inch on the lower sur- 
face, and perhaps 3,000 on the upper; and they are so re- 
markably large that they may be discerned by a simple lens of 
an inch focus. 

172. Venation, the veining of leaves, &c., relates to the mode 
in which the woody tissue, in the form of ribs, veins, &e., is 
distributed in the cellular. There are two principal modes, the 
parallel-veined and the reticulated or netted-veined. The former 
is especially characteristic of plants with endogenous stem and 
monocotyledonous embryo, and also of gymnospermous trees, 


FIG. 138. A highly magnified piece of the epidermis of the Garden Dalsam, with 
three stomata (after Brongniart). 

FIG. 139. Magnified view of the 10,000th part of a square inch of the epidermis of 
the lower surface of the leaf of the White Lily, with its stomata. 140. A single stoma, 
more magnified. 141. Another stoma, widely open. 


i i eel 


THEIR STRUCTURE AND FORMS. 91 


which have exogenous stems and at least dicotyledonous 
embryos. The latter prevails in ordinary plants with exogenous 
stem and dicotyledonous embryo. 

173. Parallel-veined or Neryed leaves (of which Fig. 143 is 
an illustration) have a framework of simple ribs (called by the 
earlier botanists zerves, 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, 


142 143 


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 flabelliform 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 (4), petiole or leaf-stallk (), and stipules (sf). 
FIG. 143. Parallel-veined leaf of the Lily of the Valley, Convallaria majalis. 


92 MORPHOLOGY OF LEAVES. 


Pasal-nerved, that is, with the nerves all springing from the 
base of the leaf, and 

Costal-nerved, 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 ; 

Curvinerved, 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- 


fined to minute and straight cross-veinlets: in many, 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 Ries (cosfe) ; the leading ramifi- 
cations, VErNs (vere) ; the smaller and the ultimate subdivisions, 
VEINLETS (venul@). All or some of the veins and veinlets are 
said to anastomose, 7. 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 etymologically rather 
misleading. More properly, it is said that the veins or veinlets 
form reticulations or net-work. A primary division of reticulated 
leaves, and indeed of nerved leaves also, into two classes, is 
founded upon the number of primary ribs. 

176. 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, 152-156) is called the Mrpris. 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 radiately divergent, giving 
a fan-shaped or rounded outline to the blade. In reticulated 
leaves, in which the veins all spring from the ribs, the two 


= es 


THEIR STRUCTURE AND FORMS. 93 


classes into which they divide are the pinnately veined and the 
palmately veined. 

177. Pinnately or Feather-veined (or Penninerved) leaves 
are accordingly those of which the veins and their subdivisions 
are side branches of a*ingle 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, &e.) 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 ¢riple-ribbed or 
tripli-nerved. ‘The appearance of a second pair of such strength 
ened veins makes the venation guintupli-ribbed ov quintupli-nerved. 


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 midrib, and each with its system 
of veins which ramify and form meshes in the interspaces. Here 
the whole woody portion of the leaf divides equally into a num- 
ber of parts upon leaving the petiole or entering the blade. The 
ribs there commonly diverge more or less in a palmate or digitate 
manner (7. e. like outspread fingers of the hand, or the claws of a 
bird, or like radii of more or less of a circle) : so, in the corre- 
lation df outline and venation, this class of veining goes with 


roundish circumscription. This is not so true, however, in a 
special case, viz., where the ribs, however divergent below, curve 
forward and all run to the apex of the blade, thus imitating the 
parallel-veined system, as in Rhexia and generally in the family 
of which that genus is the single northern representative.’ 

180. Forms as to Outline, &e. DeCandolle conceived the shape 
of leaves (both the general circumscription and the special con- 
figuration) to depend on the distribution of the ribs and veins, 
and quantity of the parenchyma in which these were outspread, 
—-a too mechanical view, and not conformable to the history of 
development. This proves that the framework is adapted to the 
parenchyma, which grows and shapes the organ in its own way, 
rather than the parenchyma to it. It were better to say that the 


1 In Linnean terminology, palmate and digitate referred to particular out- 
line only, and were separately used to denote extent of division, — palmate, not 
divided down to the petiole, digitate, when divided, like the claws of a bird, 
quite down the base. DeCandolle generalized the use of the former term, 
and ever since the two have been used interchangeably. 


FIG. 158-166. Various forms of simple, chiefly palmately veined leaves. 


THEIR STRUCTURE AND FORMS. 95 


two elements of the structure are correlated. Descriptive terms 
applied. to leaves are equally applicable to all expanded organs 
or parts, and indeed to all outlines. Some leading forms are 
here enumerated ; and all are defined in the Glossary. 

181. As to general Circumscription, proceeding from narrower 
to broader shapes, and then to those with either narrowed or 
notched base, leaf-blades are 


167 168 


170 


172 


Linear, when narrow, several times longer than wide, and of 
about the same breadth throughout. (Fig. 167.) 

Lanceolate, or Lance-shaped, when several times longer than 
wide, and tapering upwards (Fig. 153, 168), or tapering both 
upward and downward. 

Oblong, when nearly twice or thrice as long as broad. (Fig. 169.) 

Elliptical, oblong with a flowing outline, the two ends alike 
in width. (Fig. 170.) 

Oval, the same as broadly elliptical, or elliptical with the 


breadth considerably more than half the length. 
Ovate, when the outline is like a section of a hen’s-egg length- 
wise, the broader end being downward. (Fig. 171, 155.) 
Obovate, inversely ovate, or 
ovate with the narrower end 
toward the base, the broader 
Cuneate, or Cuneiform, that is, We a eed i oad cee and 
tapering by straight lines to an acute base. (Fig. 176, 148.) 
Spatulate, rounded above, long and narrow below, like a 
Oblanceolate, inverted lance-shaped, 7. e. such a lanceolate 
leaf as that of Fig. 168, but with the more tapering end at base, 
as in Fig. 173. To those who restrict the term lanceolate to the 


Orbicular, or Rotund, circu- 
lar in outline, or nearly so. 
(Fig. 160.) 
upward. (Fig. 175, 145.) eee oe 
spatula. (Fig. 174, 147.) 
sense of a narrow leaf tapering equally in both directions, the 


FIG. 167-176. Outlines of various simple leayes. 


G6 MORPHOLOGY OF LEAVES. 


term oblanceolate is superfluous. The following terms desig- 
nate leaves with a notched instead of narrowed base. 

Cordate, or Heart-shaped, when a leaf of an ovate form, or 
something like it, has the outline of its rounded base turned in 
(forming a notch or sinus) where the stalk is attached, as in 
Fig. 172, 151. Also Fig. 149, Pontederia, a leaf of the parallel- 
veined class. 

Reniform, or Kidney-shaped, like the last, only rounder and 
broader than long. (Fig. 158.) 

Auriculate, or Eared, having a pair of small and blunt pro- 
jections, or ears, at the base, as in Magnolia Fraseri, Fig. 178. 

Sagittate, 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.) 

flastate, or Halberd- 
shaped, when such 
lobes at the base point outwards, giving the leaf the shape of the 
halberd of the olden time, as in Polygonum arifolium (Fig. 179) 
and Sorrel, Fig. 163. 

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 Brasenia, of Nelumbium, and of Hydro- 
cotyle umbellata are marked examples. The anomaly is mor- 
phologically explained by a comparison with deeply cordate or 
reniform leaves haying a narrow sinus, such as those of Nym- 
pheea or Water Lily, and by supposing a union of the approxi- 
mated edges of the sinus. Fig. 159 and 160, from two species 
of Hydrocotyle, one with open and the other with closed sinus 
obliterated by the union, illustrate this. 

185. As to Extremity, 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.) 


FIG. 177-179. Sagittate, auriculate, and hastate leaves. 


i ee ee 


THEIR STRUCTURE AND FORMS. aK 


Arute, 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. 

Retuse, 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, 7. e. like 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. 188. 

Cuspidate, tipped with a sharp and rigid point, as in Fig. 187. 


184. As to Margin or special Outline, the terminology proceeds 
upon the convenient supposition of a blade with quite entire margin, 
but subject to incisions, which give rise to notches or clefts, if we 
regard the sinuses ; or to teeth, lobes, segments, &c., if we regard 
the salient portions between the sinuses. The ribs, or the stronger 
veins, &c., commonly terminate in the teeth or lobes ; but in Cicuta 
maculata, and in a few other cases, they run to the notches. 

185. Dentation relates to mere marginal incision, not extend- 
ing deeply into the blade. The blade is said to be 

Entire} when 
the margin is com- 
pletely filled out 
to an even line, as 
in Fig. 173-179. 

Serrate, when 
with small and | 
sharp teeth direct-} >} 
ed forward, a 
the teeth ofa saw, \ 
as in Fig. 189. Se 

Serrulate is the ‘159 <8 191 
diminutive of serrate, and is eaten to minutely serrate. 


1 Integerrimus-a-um, or quite entire, is the term in Latin terminology. Inte 
ger Means undivided or not lobed. 
FIG, 180-188. Terminations of leaves. 
FIG. 189-194. Dentation of pinnately veined leaves. 


7 


98 MORPHOLOGY OF LEAVES. 


Dentate, or Toothed, a general term for toothing, specially 
applied to the case of salient teeth which are not directed for- 
ward or towards the apex of the blade, Fig. 190. 

Crenate, or Scalloped, the same as dentate or serrate, but with 
teeth much rounded, Fig. 191. 

Repand, or Undulate, when the margin is a wavy line, bending 
slightly inward and outward, Fig. 192. 

Sinuate, when this wavy line is stronger or distinctly sinuous, 
as in Fig. 193. 

Incised, when cut by sharp and irregular incisions more or less 
deeply, Fig. 194. This is intermediate between dentation and 

186. Lobation or Segmentation. When the blade is more 
deeply penetrated by incisions from the margin, that is, when 
the spaces between the ribs or principal veins are not filled to 
near the general outline, it is said to be Jobed, cleft, parted, or 
divided, according to the degree of separation ; and the portions 
are called lobes, segments, divisions, &e. The most general name 
for such parts of any simple blade is that of /vbes. More par- 
ticularly a leaf-blade, or other body, is said to be 

Lobed, when the division extends not more than half way down, 
and either the sinuses or the lobes are rounded ; 

Cleft, when the division is half way down or more, and the 
lobes or sinuses narrow or acute ; 

Parted, when the divisions reach almost, but not quite, to the 
base or the midrib ; 

Divided, when they sever the blade into distinct parts, which 
makes the leaf compound. (193.) 

i187. Lope is the common name of one of the parts of a simple 
blade, especially when there is only one order of incision. But 


when there are more, as when a leaf is divided or parted and— 


these primary lobes again lobed or cleft, the lobes of first order 
are commonly called SeGMENTs (sometimes divisions or partitions), 
and the parts of these, Lobes. Or the lobes may be designated 
as primary, secondary, tertiary, &c. Ultimate portions or small 
lobes may be called Lobules or Lobelets. Also the portions of a 
quite divided blade take the name of Leaflets. By proper selec- 
tion of terms, the degree of division or lobing may thus be 
expressed in a single word. : 

188. As to Number of parts, this may be tersely expressed by 
combination with the adjective term applicable to the degree ; as, 
Two-lobed, Three-lobed, Five-lobed, Many-lobed, &c. ; or Two—Five- 
cleft, Many-cleft, &c., in Latin form Bifid, Trifid, Multifid, &e. ; 
Two-five-parted, &c., according to the number of divisions 
which extend almost to the base or axis; Two—Five-divided (in 


————— 


THEIR STRUCTURE AND FORMS, 99 


Latin form Bisected, Trisected, &c.), when there are two or three 
or more complete divisions of the blade. 

189. As to Arrangement of parts, this may be simply and best 
expressed by taking into account the nature of the venation or 
the distribution of the ribs, &e., which controls or is co-ordinated 
with the disposition of the lobes. Pinnately veined leaves, 
when lobed, must needs have the incisions directed to the mid- 
rib; palmately-veined or radiated, to the apex of the petiole. 
The lobes or divisions of the first will be pinnately, of the second 


palmately disposed. Accordingly, the three leaves of as many 
species of Oak, Fig. 195, 196, and 197, represent respectively a 
pinnately lobed, pinnately cleft, and pinnately parted leaf, while the 
accompanying leaf of Celandine, Fig. 198, is pinnately divided. 
The first three, however, when the degree of incision 1s not par- 
ticularly in question, usually pass under the common term of 
pinnatifid, Fig. 195 moderately, Fig. 197 deeply. The number 
of lobes, when definitely marked, may come into the descriptive 
phrase, as pinnately 7-lobed, pinnately 7-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, tripertite, and trisect or trisected 
leaf. Fig. 166 is a palmately 5-parted leaf; Fig. 164, palmately 


FIG. 195-198. Pinnately lobed, cleft, parted, and divided leaves. 
FIG, 199-202. Palmately 3-lobed, cleft, parted, and divided leaves. 


100 MORPHOLOGY OF LEAVES. 


multifid, &c. Fig. 162, a leaf of Dragon Arum, is palmately 
9-parted. But, as the lateral sinuses are not so deep as the 
others, the leaf is said to be pedately parted, or pedate, in the 
early terminology. 

191. Moreover, as the lobes or divisions of a leaf may be 
again similarly lobed or parted, &c., this composition may be 
indicated by the prefix twice, thrice, &¢., as twice pinnatifid or 
bipinnatifid, thrice pinnately parted, thrice palmately 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 ; for the one may pass variously into the other. Simple 
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. 

193. Compound Leaves are those which have from two to 
many distinct blades, on a common leafstalk. These blades, 
called LearLets, may be sessile on the common leafstalk, or they 
may have leafstalks of theirown. 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 merely parted, 
as in the upper portion of Fig. 198. Such leaves are so inter- 
mediate between simple and compound that it becomes indiffer- 
ent, 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. 

194. 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 types. A _pin- 
nately veined simple leaf is the homologue of one kind of com- 
pound leaf; a radiately veined leaf, of the other. That is, 
compound leaves are either pinnate or palmate. 

195. Pinnate Leaves (Fig. 205-205) are those in which the 
leaflets are arranged along the sides of a petiole, or rather of its 


THEIR STRUCTURE AND FORMS. 101 


prolongation, the Ruacuts, which answers to the midrib of a pin- 
nately veined simple leaf. There are three principal sorts, and 
some subordinate ones. That is, a pinnate leaf may be 


Impari-pinnate, or pinnate with an odd leaflet, 7. e. a terminal 
one, as in Fig. 203; and this is the commoner case. 

Cirrhiferous Pinnate, or pinnate with a tendril (Fig. 204), as 
in the proper Pea tribe and Bignonia. Here either the termi- 
nal leaflet only, or the upper lateral leaflets also, are replaced 
by tendrils. 

Pari-pinnate, or Abruptly Pinnate, destitute of a terminal leaflet 
or of any thing answering to it, as in Fig. 205. 

Interruptedly Pinnate denotes merely a striking inequality of 
size among the leaflets : Lyrately Pinnate, one in which the termi- 
nal leaflet is largest and the lower small. 

196. Palmate or Digitate Leaves (Fig. 206, 93) are those in 
which the leaflets all stand on the 
summit of the petiole. Digitate 
(fingered) was the old name, when 
the term palmate was restricted to 
a simple but palmately lobed leaf 
of this type. But since the time 
of DeCandolle the two names have 
been used interchangeably. Pal- 
mate leaves have no primary dis- 
tinction into sorts, except as to 
the number of leaflets. These can 
never be very numerous; but there are fully a dozen in some 


FIG. 203. An impari-pinnate or odd pinnate leaf. 204. Pinnate with a tendril 
205. Abruptly pinnate leaf of a Cassia. 
FIG. 206. Palmately or digitately 5-foliolate leaf of a Buckeye, Zsculus. 


102 MORPHOLOGY OF LEAVES. 


Lupines. More commonly there are only five to nine, or only 
three, rarely two, or even a single one. 

197. Number of leaflets may be indicated by an adjective 
expression composed of the proper Latin numeral prefixed to 
foliolate (Foliolum, diminutive of foliwm, answering to leaflet). 
Thus, bifoliolate, of two leaflets ; trifoliolate, of three leaflets; 
guadrifoliolate, of four; quinquefoliolate, of five: plurifoliolate, or 
multifoliolate, of several or numerous leaflets, &c. These terms 
are still more descriptive when accompanied by the word pin- 
nately or palmately, indicative of the kind of compound leaf; as, 
palmately or digitately trifoliolate (common Clover-leaf, Fig. 211), 
or 5-foliolate, as in Buckeye (Fig. 206), and so on. Also, pinnately 
16-folvolate, as in Fig. 205, or 17%-foliolate, as in Fig. 203; 
pinnately trifoliolate,as in Phaseolus, and in the low Hop-Clover, 
Trifolium procumbens.' 

198. But, in either class of compound leaves, the leaflets may 
be reduced to a minimum number. A pinnately trifoliolate leaf 
is one of the impari-pinnate kind reduced to three leaflets, to one 
pair and the odd one; and this is distinguished from a palmately 
trifoliolate leaf by the attachment of the pair at some distance 
below the apex of the petiole, and by the articulation above this, 
which marks the junction of the terminal leaflet’s petiole (or its 
base, if sessile) with the rhachis or common petiole. 

199. Unifoliolate compound leaves (by no means a direct con- 
tradiction in terms) are by this articulation distin- 
guished from simple leaves which they simulate. 
See the leaf of the common Barberry, Fig. 207. 
In other species, of the Mahonia section, the leaves 
are all pinnately 5-9-foliolate, with well-developed 
common petiole: in the true Berberis, they are all 
thus reduced to the terminal and long-petiolulate 
leaflet, on an almost obsolete petiole. Orange and 
Lemon leaves are in similar case, but with the joint 
close to the blade. A comparison with near rela- 
tives shows that these are also unifoliolate leaves 
of the pinnate kind; though this could not be ascertained by 
inspection. 

200. Decompound or Twice and Thrice Compound Leaves. These 
are to once pinnate or once palmate leaves what the latter are to 


1 In pinnate leaves, each leaflet usually has its opposite fellow, and the 
number may be indicated by the pairs, as unijugate, bijugate, trijugate, and 
plurijugate, according to the number of juga, or pairs. 

FIG. 207. Unifoliolate leaf of Berberis vulgaris, with partial petiole articulated to 
the extremely short true petiole. 


THEIR STRUCTURE AND FORMS. 103 


simple leaves. As leaflets may be toothed, lobed, or parted, so 
what answers to a single leaflet may appear as leaflets of a second, 
or again of a third, or even of a fourth order. Decompound is a 
good general name for all more than once compounded leaves ; 


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but the name has been applied rather to irregularly many-times 
parted or dissected leaves (such as those of Dicentra), or to 
those more than thrice compounded. Of regularly twice or 
thrice compound leaves, the commonest are the 

Bipinnate or Twice Pinnate, of ordinary 


occurrence in the Mimoseous and Cvesalpi- ; 
neous, but not in the Papilionaceous, Legu- a 
minose. Fig. 208 represents a bipinnate leaf if y 


of the Honey Locust (Gleditschia), with the 
variation (common with that tree) that some 
of the partial petioles, in this figure only the 
lowest, bears a single leaflet, while the others 7 ‘ 
are extended into secondary rhachises fur- } 


nished with numer- ue 
ous leaflets, mostly cS e pie 
j : > pin- <—Sdioapsse /) ; 
in the abruptly pin-  <Sy ie: 5 


iN 
te style. On th = 14 = WOH H i \ 
ae Bee the are = (I Uijz= = i H y 
leaves, which are iy! // “is | ; : U 
lustered hort <7 =: J 
clustered on sho =i lS 


spurs, are simply 
pinnate. The large 
leaves of Gymnocladus are similarly and abruptly bipinnate, 


FIG. 208. A bipinnate and multifoliolate leaf of Gleditschia or Honey Locust. 
FIG. 299. Bipinnate leaves of Sensitive Plant, Mimosa pudica, with approximate 
pinne. 


104 MORPHOLOGY OF LEAVES 


except at the base, which is simply pinnate or with one or two 
pairs of simple leaflets. 

Tripinnate or Thrice Pinnate leaves of a regular sort are rare ; 
but, with some irregularity, they occur in many species, as in 
Aralia, &c. This extent of division, and even much greater, is 
common in Ferns. 

Digitate-Pinnate is where the primary division of the petiole is 
on the palmate or digitate plan; the secondary, on the pinnate. 
This seems to be the case in the Sensitive Plant, Mimosa pudica, 
Fig. 209. But the leaf is here truly bipinnate with the primary 
divisions very crowded at the apex of petiole. 

Conjugate-Pinnate 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 Palmately Decompound in a nearly reel way 
is not an uncommon case. 
Usually, the petiole is succes- 
sively three-forked, as in Fig. 
210, when the leaf is said to 
be biternate (twice ternate), 
triternate (thrice ternate), or 
quadriternate (four times ter- 
nate), &c., according to the 
number of times it divides, or 
2-3—4-times ternately compound. 

an The ultimate divisions in such 
cases of threes are commonly of the pinnately trifoliolate type. 

201. Pinne is a convenient name for the partial petioles of a 
bipinnate leaf, taken together with the leaflets that belong to 
them. Thus, the Sensitive Plant, Fig. 209, has four pinne, or 
two pairs; the Honey Locust, Fig. 208, a greater number. 
When such leaves are still further compounded, the pinn of 
higher order, or the ultimate ones, take the diminutive term of 
PrxnuL& or Pixnutes. ‘The blades these bear are the Leaflets. 

202. The Petiole or Leafstalk is a comparatively unessential 
part of the leaf. It is often wanting (then the blade is sess¢le) ; 
it may be 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 
the blade, so that the slightest breath of air puts the leaves in 
motion. Sometimes it is much dilated and membranaceous at 


FIG. 210. Quadri-ternately compound or ternately decompound leaf of Thalictrum 
Cornuti. 


THELR STRUCTURE AND FORMS. 105 


base, as in many umbelliferous plants; sometimes it forms a 
sheath, occasionally it is bordered with appendages, &c.  Peti- 
oles may 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 ramified in the blade. The ends of these threads 
are apparent on the base of the leafstalk when it falls off, and on 
the scar left on the stem, as so many round dots (Fig. 81, 85, 
91), of a uniform number and arrangement in each species. 

203. Partial Petioles are the divisions of the petiole in a 
compound leaf. The footstalk of a leaflet takes the diminutive 
name of PETIOLULE. 

204. Stipules (157) are lateral appendages, one each side of 
the base of the petiole, sometimes free 
from it and from each other (Fig. 
142), sometimes attached by one 
edge to its base (Fig. 211), some- 
times united with each other into a 4 
single body (Fig. 212) in various 
ways or degrees. In the latter case, 
they usually appear to be within the 
base of the leaf or leafstalk; or, as 
in the Plane-tree, they may be joined 
into one over against the leaf, as 
if opposite to it, but their normal 
position is supposed to be lateral or 
marginal to the petiole. Sometimes 
they are foliaceous in appearance and 
in function; sometimes they are dry 
and colorless or scale-like, reduced to 
mere epidermal tissue, and evidently 
functionless ; sometimes (as in Mag- 
nolia, Fig. 81, Fig-tree, and Beech), 
they serve as bud-scales, and fall when 
the leaves develop ; sometimes they are 
reduced to a mere bristle, or take the a a 
form of a spine, as in the Locust (Robinia). Between salient 
expansions or wing-like margins of the base of the petiole, 
such as those of the Saxifrage tribe, and stipules adnate to the 
margins of the petiole, as in most Rosacew, there is no clear 
limitation. But presence or absence of stipules generally runs 


FIG, 211. Clover-leaf, with adnate stipules. 212. Ochreate stipules (ochrea or ocrea! 
of Polygonum orientale, sheathing the stem for some distance, and ending in a spread- 
ing border. 


106 MORPHOLOGY OF LEAVES. 


through a naturai order. Yet what are called stipules in one order 
may pass for expansions or appendages of the petiole in another. 
In Spergularia, some stipules are connate around the base of the 
pair of leaves, including them as well as the stem in the sheath." 

205. Stipules, which are normally a pair, may unite into one 
body, either adnate to the inner face of the leaf, as in some 
species of Potamogeton, or united opposite the leaf, as in Plane- 
tree, or united znfer se in a sheath, as in Polygonum. Also when 
the leaves are opposite and the stipules thus brought into prox- 
imity, the adjacent half stipules of the two leaves may coalesce, 
and present the appearance of only two stipules to two leaves, 
as in many Rubiacez. A notch or fork at the apex often 
indicates the composition. 

206. Sheathing stipules, like those of Polygonum (Fig. 212), 
are said to be oehreate, or (better) ocreate ; the sheath, thus 
likened to a leggin or the leg of a boot, is an OcureEa, as written 
by Willdenow, or better Ocrra. 

207. The Licute of Grasses (Fig. 150) is seemingly a thin and 
scarious 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 Leguminosee, e. g. 
in the Phaseolex, in Wistaria, Locust. &c.; also in Staphylea. 
They 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 st/pulate, so leaflets 
with stipels are st/pellate. 

209. Some unusual modifications of leaves as foliage. In 
leaves as illustrated thus far, it is the lamina or blade which is 
expanded to do the work of foliage; which is expanded hori- 
zontally, so as to present upper and under surfaces, one to the 
sky, the other to the ground; which is bilaterally symmetrical 
or substantially so, the two lateral halves being nearly if not 
quite alike ; and which is affixed to the stem at the basal margin, 
or some part of it, with or without a petiole. Various deviations 
or apparent deviations from this pattern occur. Some of them are 
of comparatively small account and simple explanation, such as 

216. Inequilateral Leaves, being unsymmetrical by the much 
greater development of one side. This is illustrated in the 
whole genus Begonia (as in Fig. 161), consisting of many spe- 


! As pointed out by Prof. A. Dickson, in Nature, xviii. 507. 


THEIR STRUCTURE AND FORMS. 107 


cies, some of which are moderately, and most of them strikingly, 
oblique in this way. Elm-leaves, and the like, are more or less 
inzequilateral at the base. 

211. Connate and Perfoliate Leaves. These are explained by 
the union of con- 
tiguous leaf-edges. 
Peltate leaves, to 
which a paragraph 
has already been 
given (182), come 
under the same 
head; the seeming 
attachment of the 
petiole to the lower 
face of the blade 
being the result of 
a congenital union 
of the edges of the 
sinus. In a sessile ? 
leaf, when such a 213 a 
union takes place, it surrounds and encloses that portion of the 
stem ; which is thus perfoliuate. (Fig. : 
213, 214.) It is the stem which is 
literally perfoliate, ¢.e. which seem- 
ingly passes through the leaf; but it 
is customary, though etymologically 
absurd, to call this a perfoliate leaf! 
Uvularia perfoliata (Fig. 213), in the 
later growth of the season, reveals the 
explanation of the perfoliation: the 
base of the lower leaves conspicuously 
surrounds and encloses the stem: that 
of the upper is merely cordate and 
clasping ; the uppermost simply ses- 
sile by a rounded base. Baptisia 
perfoliata (Fig. 214) is a more 
strongly marked case of perfoliation. 
But there are good morphological 
reasons for inferring that this seemingly simple leaf consists of 
a pair of stipules and a leaflet combined. An occasional mon- 
strosity verifies this supposition. 


WM 
KW \ a 


FIG. 213. Leafy branch of Uvularia perfoliata. 

FIG. 214. Leafy and flowering branch of Baptisia perfoliata. 

FIG. 215. Lonicera flava, a wild Honeysuckle, connate-perfoliate as to the upper 
leaves. 


108 MORPHOLOGY OF LEAVES. 


212. When leaves are opposite, the perfoliation (such as that 
of Honeysuckles, Fig. 215) is obviously the result of a congeni- 
tal union of the bases of the pair by their contiguous edges. 
Leaves connate in this way by narrow bases are not rare nor 
remarkable ; but when the two are thus coalescent into one broad 
foliaceous body, giving this appearance of perfoliation, the term 
connate-perfoliate is used to express it. 

213. 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 Myrtacez, &¢., and occur with 
less constancy in the Californian 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.! In strongly marked 
cases, or in most of them, the organization of 
the epidermis and superficial parenchyma and 
the distribution of the stomata are the same on 
both faces. 

214. Equitant Leaves are vertical on a different L\'t 
plan. They are conduplicate, ¢.e. are folded 6 * ¥ 

&> 


UMN 


E> yy 


217 26 
together lengthwise on their middle, the upper surface thus con- 
cealed within, the outer alone presented to the air and light. 


atta as 
i 


1 Silphium laciniatum, the so-called Compass Plant, and (hardly less so) 
S. terebinthinaceum, are 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 pinnately parted blade occasionally makes a farther twist, so as 
to bring the upper part into a plane at right angles to the lower. The 
blades place themselves in various directions as respects the cardinal points ; 
but on the prairies the greater number affect a north and south direction of 
their edges,—a peculiarity first pointed out, in the year 1842, by General 
B. Alvord, U.S. A. 


FIG. 216. Equitant erect leaves of Iris, with the rootstock. 
FIG. 217. A section across these leaves at the base, showing the equitant character. 


THEIR STRUCTURE AND FORMS. 109 


Being two-ranked and closely crowded, the outer ones at their 
base fold over or bestride the inner (as shown in the sectional 
diagram, Fig. 217), whence the name of eguitant. Above, the 
contiguous halves of the inner face congenitally cohere, and so 
produce the sword-shaped or linear vertical blade which is 
characteristic of Iris (Fig. 216) and the Iris family. In most 
there is a farther complication, of an excep- 
tional kind, viz. the development backwards 
of a portion of blade from the midrib, often 
forming most of the upper part of such leaves, 
which therefore may really be said to develop 
in the vertical plane. 

215. Leaves with no distinction of Parts, 7. e. 
of blade and petiole. ‘This is the case in Iris 
(Fig. 216), Daffodil, the Onion, and perhaps 
of most parallel-veined leaves of Endogens. 
Those expanded in the horizontal plane may 
however be regarded as sessile blades: those 
which are not expanded, but filiform, or needle- 
shaped (acicular), or awl-shaped (subulate), may 
be regarded either as homologous with petioles, 
or as unexpanded blades, which amounts nearly a4 
to the same thing where there is no trace of a petiole at base. 
Under this head may be ranked the leaves of Pines (Fig. 248) ; 
also both the subulate and the 
scale-shaped and adnate leaves 
of Arbor Vite, Red Cedar (Juni- 
perus Virginiana), and other trees G 
of the Cypress tribe. (Fig. 218.) ~~ 

216. Stipules serving for Blade. 
Lathyrus Aphaca is a good in- 
stance of this (Fig. 219); the 
petiole becoming a tendril, the 
leaflets which its relatives bear 
being wholly wanting, the ample 
foliaceous stipules assume the 
appearance of leaves. In some on 
other species of Lathyrus, and in the Pea, equally large stipules 
share with the pair or pairs of leaflets in the functions of foliage. 
On morphological evidence, we judge that the singular leaves of 


FIG. 218. A twig of Arbor Vite, with both awl-shaped and scale-shaped leaves. 

FIG. 219. Lathyrus Aphaca: portion of stem, bearing a single leaf, which consists 
of a pair of foliaceous stipules, and a petiole in the form of a tendril; in its axil a 
flower-stalk. 


110 MORPHOLOGY OF LEAVES. 


Baptisia perfoliata, shown in Fig. 214, are not simple blades, 
but each a pair of stipules, with or without a terminal leaflet, all 
completely confluent into one body. The related species of the 
genus have trifoliolate leaves and foliaceous stipules ; hence these 
simple leaves without stipules are best explained in this way. 

217. Phyllodia, or Petioles serving for Blade. Sometimes the 
petiole develops foliaceous margins, or wings, as in the Bitter 
Orange and in Rhus copallina. These are efficient as foliage in 
proportion to their size. These are not to be confounded with the 
case in which a petiole specially develops as a blade-like organ, - 
which usurps the office of foliage. A petiole-blade of this kind 
is named a Puyttopium. Occurring only in Exogens, phylodia 
are generally distinguished from true blades by the parallel 
venation, and always by their normally vertical dilatation ; 7. e. 
they, without a twist, present their edges instead of their faces to 
the earth and sky. The common and most familiar phyllodia 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 phyllodia. So likewise do our tubular or trumpet- 
leaved species of Sarracenia in that portion of the foliage which 
develops the pitcher imperfectly, or not at all. Indeed, all 
Sarracenia-leaves are phyllodia with the back in most of them 
hollowed out into a tube or pitcher; and the terminal hood 
answers to the blade. 


§ 3. Leaves servine 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 
for a 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 

219. 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. 


eet 


subject — which has of late become highly interesting — belong 
to physiology, and therefore to the following volume, to which 


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, always 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 nndergoing macer- 


ation. InS. 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 Heliamphora; 221. of Sarracenia purpuren; 


222. of Nepenthes. 


223. A phyllodium of a New Holland Acacia. 224. The same, bearing a reduced com- 


pound blade. 


FIG. 225. Pitcher-leaves of Sarracenia purpurea; one of them with the upper part 


cut away. 


ya Wy MORPHOLOGY OF LEAVES. 


as to mostly exclude the rain; in 8. psittacina (Fig. 227) the 
inflexed and 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 
sweetish secretion around or at some part of the 
orifice.t Few that have entered ever escape ; 
most are decomposed at the bottom of the cavity. 
In Darlingtonia Californica (Fig. 228), the Cali- 
fornian representative of Sarracenia, the inflated 
hood guards against all access of rain, while the 
orifice is freely open to flying insects from be- 
neath; and a singular two-forked appendage, like 
to a fish-tail (probably the homologue 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.? Fig. 220 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- 
226 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-fitting lid, 
hinged by one edge, Fig. 229. « 
The particular morphology of the parts is not well made out. 


1 This sweet secretion, which at times is very obvious in the southern 
species, has also been detected by Mr. Edward Burgess 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. 

2 This trail was discovered by Dr. J. H. Mellichamp, of South Carolina. 
See Proce. Am. Association for Advancement of Science, xxiii. 113 (1871). 


FIG. 226. Pitcher of Sarracenia variolaris. 227. Same of S. psittacina. 


LEAVES SERVING SPECIAL OFFICES. 143 


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 may be 
regarded as a prolonged 
extension of the midrib 
of the blade, and the 
pitcher, with its hinged 
lid, as a peculiar development from. as 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.’ 

222. The aquatic sacs of Utricularia or Bladder- 
wort are diminutive ascidia, always under water, 228 
and with lid opening inward, like a valve, preventing the exit 
of minute animals entrapped therein.*? 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. D. Hooker, and announced in his address, 
as President of the British Association for the Advancement of Science, at 
Edinburgh, 1871. 

2 Darwin, Insectivorous Plants, 395. Cohn, Beitrige zur Biologie der 
Pflanzen, 1875. Mrs. Treat, in The Tribune, New York, September, 1874, 
and Gard. Chron. 1875, 303. 


FIG. 228. Pitcher of Darlingtonia Californica. 229. Pitcher of Cephalotus follicu- 
laris, with lid open. 


114 MORPHOLOGY OF LEAVES. 


tion of petiole and blade, and the stalked glands thickly beset 
the whole upper surface of the latter. A small insect alighting 
thereon is helpless, and is soon touched by all the glands within 
reaching distance; also the blade itself commonly ineurves, 
taking part in the general movement. It has recently been 
demonstrated that the captured insect is fed 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 


A 
we 


also the power and the habit of obtaining ready-organized food 
by capture, and are benefited by it. 

224. Species of Drosera inhabit most parts of the world, and 
the genus is numerous in species. A near relative, Dionza, is 
of a single species, D. muscipula (Venus’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. 230, 231.) If this 


FIG. 230. A plant of Dionwa muscipula, reduced in size. 231 Three of the leaves, 
of almost the natural 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. de 5) 


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 ready for another capture. For 
references to the now copious literature of this whole subject, and 
for its physiological 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 A pT 
same time are subserving the purpose of foliage. / //// f\\ 
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. 252, a leaf from 
the bulb of White Lily, the base of which forms 
one of the bulb-scales, is an instance of the kind.! 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, are turned to this 
account. (21-37, &c.) After or 
while discharging this special duty, 
the cotyledons may fulfil their gen- 
eral office, by serving as foliage (as 
in Maples, Fig. 8, and Pumpkins, 
Fig. 47) ; or, through various inter- 
mediate conditions, they may be 
wholly devoted to storage, as in 
the Pea, Oak, Horsechestnut, &e. 
(Fig. 37-43.) 

226. Leaves as Bulb-seales, how- 
eyer, are for the most part wholly 
applied to this use, being leaves 
reduced to short scales or to 
concentric coats, and thickened 
throughout by nutritive deposit. 
The accumulation of such leaves 
forms the mass of the bulb, as of 
the Lily, Fig. 118, Onion, Fig. 
113, &e., also of bulblets. (120.) 

227. Leaves as Bud-seales, being 
for protection of nascent parts, have 
been explained under buds. (70.) 
The evidence of foliar nature af- 
forded by transition is well exhib- 
ited by the Sweet Buckeye, although 
the whole series of gradations, from 
bud-scales to compound leaves, 
is seldom seen united in one bud, 
as in Fig. 233. In this case, the 
bud-seales are homologous with 
petioles. In Magnolia, they consist 
of stipules (Fig. 81, 82): in the 
Lilac, they are homologous with leaf-blades. The two pairs of 
bud-seales which subtend and protect through winter the nascent 
head of flowers of Cornus florida are morphologically the apex of 


centrated condition, in the form of a solid grain, which remains for next 
year’s use, the whole leaf except this thickened base dying away at the close 
of the short season’s growth. 

FIG 233. Leaves of a developing bud of the Low Sweet Buckeye (#sculus parvi- 
flora), showing a nearly complete set of gradations from a scale to a compound leaf ot 
five leaflets. 


LEAVES SERVING SPECIAL OFFICES. 417 


blades. When the blossoms develop in spring, these scales grow 
from beneath, greatly expand, and become obovate or obcordate 
petaloid leaves, the brown terminal notch 
of which is the bud-scale, which was un- 
able to take part in the vernab growth. 

227%. Leaves as Spines. All gradations 
may be found between spiny-toothed leaves 
(as in Holly), in which teeth are pointed and 
indurated, and leaves which are completely 
contracted into a simple or multiple spine. 
Indeed, such a transition is seen in the Bar- 
berry, Fig. 254. The foliar nature of such 
spines is manifest from their position, sub- 
tending a bud from which the foliage of the 
season proceeds, and themselves not sub- 
tended by any organ. In some Astragali, 
the petiole of a pinnate leaf indurates into 
a slender spine and persists, the leaflets 
early falling. The spine in Fouquiera is a 
portion of the lower side of the petiole or 
midrib, indurated and persistent, the rest 
of the leaf separating by splitting when it 
has served its office.! 

228. Leaves adapted to Climbing. Some plants climb by the 
action of the stem or of certain branches specially ue to 
this purpose (99): others gain the needful 
support by means of their leaves (101) ; some- 
times by an incurvation of the tips, either of 
a simple blade as in Gloriosa, or small partial 
blades, as in Adlumia, and often in Clematis, 
thereby grappling the support ; sometimes by 
the petiole making a turn or two around a 
support (as in Maurandia, climbing Antirrhi- 
nums, Rhodochiton, and Solanum jasminoides, § 
Fig. 235) ; sometimes by the transformation f4 
of one or more leaflets of a compound leaf 7 ¥ 
into tendrils, as in the Pea and Vetch (Fig. 
204) ; sometimes by the suppression of all the 
leaflets and the conversion of the whole petiole into a tendril, as 
in Lathyrus Aphaca (Fig. 219) ; and perhaps by the conversion 


234 


1 Described in Plante Wrightiane, ii. 65. 
FIG. 234. A vernal shoot 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 Cucurbitaceze are peculiar and ambiguous, on account of their 
lateral and extra-axillary position and the manner in which the 
compound ones develop their branches. But they are doubtless 
partly if not wholly foliar.’ 

229. Petaloid Leaves, Bracts. Certain leaves, situated near to 
flowers, 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 petalord, i. e. petal-like. They 
are like petals, moreover, in one of the purposes which these sub- 
serve. (299.) Examples of these petaloid leaves are seein in the 
shrubby Mexican Euphorbia called Poinsettia, in Salvia splen- 
dens, most species of Castilleia or Painted Cup, also in the 
white hood of Calla and Richardia /thiopica (called Calla Lily), 
and in the four white leaves which subtend the flower-head of 
Cornus florida, and of the low herbaceous Cornel, C. Canaden- 
sis. (Fig. 294.) 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 Bracrs. More usually 
bracts are not petaloid, but different in size 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. 

230. 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 be preceded 
by a study of the arrangement of leaves and of blossoms. 


1 The most satisfactory interpretation may be that of Braun and Wydler, 
adopted by Eichler (Bliithendiagramme, i. 304): that the flower of Cucur- 
bita and its peduncle represent the axillary branch, the tendril by its side 
answers to one of the bractlets (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 in 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, OR LEAF—ARRANGEMENT. 119 


CHAPTER IV. 


PHYLLOTAXY, OR LEAF-ARRANGEMENT. 


Section I. THe DIstrisuTION oF LEAVES ON THE STEM. 


231. Puytiotaxy (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.! Leaves 
are arranged in a consider- 
able variety of ways, which 
all fall under two modes, the (==> 
Verticillate and the Alter-\G 
nate (13), but which may 
also be termed the Cyclical g 
and the Spiral. f 

232. Alternate leaves are 
those which stand singly, 
one after another; that is, 
with one leaf to each node i 
or borne 236 237 
on one height of stem. Verticillate leaves are 
those with two or more at the same height of 
stem, circularly encompassing it, 7. 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 
Morphologie, § 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 are 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 be kept in mind that 
“leaves opposite ” is the same as ‘* leaves in whorls of two.” 

233. The greater number of phanogamous plants (all but the 
monocotyledonous class) begin with verticillate leaves, mostly 
of the simplest kind (7. 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 
readily from the one mode to the other on the same axis, we 
may 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.! 

254. Verticillate or Cyelical 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, quaternate, quinate, 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, 7. e. they divide the circle equally. Thus, when 
only 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, 90°; when five, 72°. 

255. The characteristic of the whorls mn 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 alternate or decussate. ‘This economizes 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 the 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. 237), the suc- 
cessive pairs deeussate or cross at right angles, and so four 


1 It is readily seen that whorls may be produced by the non-development 
of the internodes between the leaves of a series of two, three, five, or more 
in alternate order. The difficulty is that the members of the next whorl do 
not follow the order that they should upon this supposition. 


DISTRIBUTION OF LEAVES ON THE STEM. 121 


straight equidistane 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.! 


236. The cases in which successive ‘S) 
pairs of leaves do not decussate at right ——— 


angles, or the members of whorls are not Fe 
exactly superposed to intervals, but as it 

were wind spirally (as in Dipsacus, many f 
Caryophyllacez, &c.), may some of them ee y 
be explained by torsion of the stem, So 


such as is very manifest in numerous in- % 


SS 


stances ; and others may be resolved into 239 
instances of alternate leaves simulating or passing into whorls 
by the non-development of internodes.? 

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. 256.) This angu- 
lar divergence (/.e. the angular distance of any two successive 
leaves) differs in the various kinds of this system of phyllotaxy, 
but is always 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, &¢., 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, 7. e. the pairs not decussating at right angles, thus 
forming double spirals. In the abnormal spruce-cones, the fractions usually 
observed are 7 or 7g, or, as expressed by Braun, (4)35 and (4)3°s. 

Braun’s mode of notation for the ordinary succession (?. e. the decussation ) 
of opposite leaves is (4)4, the 4 meaning that the two leaves of the pair are 
half the cireumference of the circle apart, the 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.g. certain species of Mesembryanthemum and Euphorbia). these 
are expressed in this notation by the formula (4)4, that is, the corresponding 
leaves of the succeeding pair diverge 180° from their predecessors. He 
recognizes also some cases of intermediate divergence; such as (4)? in the 
upper leaves of Mercurialis perennis, (}),°; on certain stems of Linaria vul- 
garis, (4)3°; exceptionally in the leaves of Epilobium angustifolium and the 
scales of Norway Spruce, (4)3 exceptionally in the scales of Norway 
Spruce. See Ordnung des Schuppen an der Tannenzapfen, 376, &c. 


FIG. 239. Ground-plan diagram of six trimerous whorls, showing their alternation. 


122 PHYLLOTAXY, OR 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 internodes 
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 spzral, 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.! 

239. Almost all 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 orrank. 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 
exactly opposite sides of the stem (as in Fig. 1); the seeond 
leaf being the farthest possible from the first, as is the third from 
the 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 180°; and 
the phyllotaxy may be represented by the fraction 4, 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 a lower and older 
one, we say that one set or spire is completed, and that this leaf is the first 
of a succeeding set or spire. 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 as a cycle. Yet it is better (with Eichler) to restrict that term, and 
the adjective cyclical, to verticillate 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 4 designates this 
arrangement. ‘Fhe 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.! 

Pentastichous, or Five-ranked, sometimes termed the guincuncial 
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 
second leaf (passing the shorter way) is 2 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 2, 


1 The line is supposed to follow the nearest way, and the divergence is 
counted as 4, 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 2. 


FIG. 240. Piece ofa stalk, with the sheathing bases of the leaves, of a Sedge-Grass 
(Carex cruscorvi), 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 
turn: as the sixth would come directly before, or within, the first. 


124 PHYLLOTAXY, OR LEAF-ARRANGEMENT. 


and the sixth is the first to come over any one below; the 
seventh comes over the second, the eighth over the third, &e. 
The leaves are thus brought into five vertical ranks; but these 
7 five leaves are laid down on two turns of the 
9 helix (the sixth beginning the second revolu- 
> | tion); the angular divergence of the leaves in 
; * order is 2, or 144°; the angular distance of 
1 © the vertical ranks, 72°. This is a very advan- 
tageous distribution for ordinary foliage on 
erect or ascending branches. Its formula is 2, 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. 
Vig. 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 Hight-ranked, a less common 
arrangement, occurs in the Holly, Aconite, 
the radical leaves of Plantago. It has the 
angular divergence of 135°, or 2 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 3. 
240. The obvious relations of the fractions 
3, 1, 2, 2, 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- 
ceeding fraction. Following these indications, 
the series may be extended to 75, #, 43, 34, &c. Now these 


v0 


10 


= 


FIG. 244. A cone of the small-fruited American Larch (Larix Americana), with 
the scales numbered, exhibiting the five-ranked arrangement. 

FIG. 245. An offset of the Houseleek, exhibiting the 5-13 arrangement; the leaves 
in sight numbered, the 14th over the first, the 19th over the 6th, &e. 

FIG. 246. Cone of White Pine (Pinus Strobus) with scales numbered from bottom, 
and some secondary spirals marked. 


DISTRIBUTION OF LEAVES ON THE STEM. #25 


eases actually occur, and ordinarily only these.’ The 4% and 
#, are not uncommon in foliage. The rosettes of the House- 
leek exhibit the 5, or thirteen-ranked arrangement, as also does 
the cone of Pinus Strobus, the 14th leaf falling over the first. 
(Fig. 246.) The 3 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 3} 
and 24 phyllotaxy may be readily made out. This actual series, 
4, 4, 2, 2, &c., answers to and may be expressed by the con- 


tinued fraction, 44 1 


iz ; 
Se ter &e.? 


1 When other instances are detected, they are found to belong to other 
series, following the same law, such as the rare one of }, 4, 3, ;%;. 

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 ? by almost exactly 7755, 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 innature. 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 
2 and 3° 3, &e., 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 34, = 0.381, and differ from & [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 & for all the values of the 
series after the first three, it should be with the understanding 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° ps = 138° 27! 41.54" 
= 120° 135° #, = 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. They 
all fall within the } and } as to amount of divergence: and they 
form a series converging to a deduced typical angle of 137° 30! 
28”, which, being irrational to the circumference, would place 
no leaf exactly 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 curvisertal, or 
actually become so; while in the lower grades they are obviously 
rectiserial. Unless. indeed, there is some torsion of the axis, 
by which the vertical ranks are rendered oblique, as is often the 
case in cones of the Norway 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 3, secures the utility of the 
theoretical angle, viz., that ‘* by 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 symmetrical 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.! 

242. It is to be noted that the distichous or 34 variety gives 
the maximum divergence, viz. 180°, and that the tristichous or 
4 gives the least, or 120°; that of the pentastichous or 2 is nearly 
the mean between the first two; that of the 2, 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, therefore placing them 
vertically farther apart; or else, as in Elms, Beeches, and the 


i ole 


Wl 
im 


1 This corresponds with Hofmeister’s general rule, that “new lateral 
members have their origin above the widest gaps 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 invokcs. 


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 2, with dimin- 
ished divergence, increase the number of ranks; the 2 and all 
beyond, with mean divergence of successive leaves, effect a more 
thorough. distribution, but with less and less angular distance 
between the vertical ranks. 

242°. The helix or primitive spiral upon which the leaves 
successively originate ascends, sometimes from left to right, 
sometimes from right to left,’ 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 already stated) the spiral 
may 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 irregularly 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, which 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 seen from without. See p. 51, foot-note. 


128 PHYLLOTAXY, OR LEAF-ARRANGEMENT. 


ranks are 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. 

245°. Take, for example, the 2 arrangement, where, as in the 
diagram annexed to Fig. 244, the primitive spiral, written on a 
plane surface, appears in the numbers, 1, 2, 3, 4, 5, 6, and so 
on: the vertical ranks thus formed are necessarily the numbers 
1-6-11; 4-9-14; 2-7-12; 5-10-15; and 3-8-13. But two 
parallel oblique ranks are equally apparent, viz. 1-3-5, which, 
if we coil the diagram, will be continued into 7-9-11-15-15 ; and 
also the 2—4—6-—8-—10 continues into 12-14, and so on, if 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 two; 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 10-15 ; 3-6-J-12, 
continued into 15; and 5-8-11-14, &c.; where again the common 
difference, 3, accords with the number of such ranks. This fixed 
relation enables us to lay down the proper numbers on the leaves, 
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. 244), 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 3-5-7, 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. 3 has already been fixed, we carry on that se- 
quence, 6-9, &c.; and on the third, where No. 45 is already 
fixed, we continue the numbering, 8-11, &c. 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 2 order. 


DISTRIBUTION 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 are 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.* Ina rosette of the leaves of Houseleek 
(Fig. 245) and a cone of Pinus Strobus (Fig. 246), 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 
ys; 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 plyllotaxy, 
leads into interesting fields. But this sketch may suffice for 
botanical uses. 

246. Relations of Whorls 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 be produced by the non-development of the 
internodes 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 
tristichous (4) arrangement would compose a trimerous whorl ; 
the five leaves of the two turns in the pentastichous (?) arrange- 
ment, a 5-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 phzenogamous plants in which the 
leaves are opposite and all in the same plane * (that is, the suc- 
cessive pairs superposed) may 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 with spiral phyllotaxy ; 
for 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.* 

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 Europzus, &c., according to Braun. See 236, note. 

2 This renders the verticillate 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 their respective utilities, and 
to have been independent determinations. For “there is no continuity or 
principle of connection between spiral arrangements and whorls ” (Chauncey 
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 five members, as in a quincuncial 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 separatcd 
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 Algze, Hepaticee, 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.’ 

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 
may 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, 7. e. two very 
short and stout branclilets, 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 bunile 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. 


132 PHYLLOTAXY, OR LEAF-ARRANGEMENT. 


Section II. Disposition or LEAVES IN THE Bup. 


249. Vernation and Estivation are terms in general use, under 
which the disposition of leaves in the bud 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), but as to 

251 the ways in which they 


249 20 
are coiled, folded, over- 
lapped, &c., either per se 
yal bast or inter se. Prefoliation 
and Prefloration are 


etymologically better 


terms, substituted by 
6O Richard.? 
250. The descriptive 
= 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 Linnean ones are 
most in use, and, though fanciful, are not misleading. In English description, 
it isas convenient and equally terse to say that the parts are imbricate, val- 
vate, &., “in the bud.” Linnezus, in the Philosophia Botanica, described 
these dispositions of leaves in the bud under the term Fo//atio, — not a happy 
name,—but did not treat of them in the flower-bud. Later, in Termini 
Botanici (Amen. Acad. vi. 1762, reprinted by Giseke in 1781), he intro- 
duced the words Vernatio and ’stivatio 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 flower-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, Linnzus fixed nearly as it now remains. That of 
leaves 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 Proteacez, 1809, 
in the Prodromus a year later, and in other publications. 

Ptyxis (the Greek name) is coming into use as a general term for the 
folding, &c., of single parts. ; 

FIG. 249-254. Linngan diagrams of sections of leaves in the bud. 249. Condupli- 


cate. 250. Plicate or plaited 251. Convolute. 252. Revolute. 253. Involute, 254. 
Circinate or Circinal j 


VERNATION, OR PRAFOLIATION. 133 


in the Philosophia Botanica of Linnzeus. They were applied 
only to foliage, but they are equally applicable to floral parts. 
Leaves, and 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 ov 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. 

Cenduplicate (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, 7. 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 Stcironema and Tremandra. 

Revolute (Fig. 252), similarly rolled backward from both 
margins, as the leaves of Azalea and Rosemary. 

Circinal ov Circinate (Fig. 254), when coiled from the apex 
downward, as the leaves of Drosera and the fronds of all the 
true Ferns. 

Oorrugate or Crumpled, as the petals of a Poppy, applies 
to the irregular crumpling of the otherwise plane corolla-leayes. 
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, 7.e. in wstivation. To this the 
following exposition is devoted, although sometimes applicable 
to leaf-buds also. 

252. The disposition of parts in wstivation, 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-leayes, and petals or corolla-leaves. See, 
if need be, Chapter VI. Sect. I. 


134 PHYLLOTAXY, OR LEAF—ARRANGEMENT. 


their phyllotaxy, and partly of the way in which they comport 
when their margins meet in growth. Those leaves which are 
within, or of higher insertion on the axis, will almost necessarily 
be enclosed or overlapped: those which are members strictly 
of the same whorl or cycle may fail to come into contact, or 
may meet without overlapping at the contiguous margins or 
apex; yet they may be 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 sestivation of 
DeCandolle), is a matter of phyllotaxy, not of wstivation. The 
latter is properly concerned only with the relations of each three 
leaves to each other.? 

253. The proper estivations 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 (est. 
aperta) and the valvate, both characterized and 
named by Brown.? Of the second, there is 
one leading kind, the zmbricate (adopted by 
Brown from Linnzeus), with subordinate modi- 
fications. Accordingly, the vestivation is 
said to be 

254. Open or Indeterminate (est. aperta), 
when the parts do not come into contact in the bud, so as to 


1 The same applies to the two sets of sepals and of petals in Barberry, in 
Menispermum, and of the petals in Poppy, &e. (359). 

2 Linneus, indeed, has, “ stivatio valvata, si petala se expansura instar 
glume graminis ponuntur,’—the name, but not the thing: the glumes of 
grasses are not valvate in the botanical sense. So the term as to its proper 
use may be said to originate with R. Brown. 

8 For a brief discussion of “Estivation and its Terminology,” see Amer. 
Jour. Sci. ser. 3, x. 339, 1875. 

As to names, it is perhaps more correct to say of the cestivation that it is 
imbricative, convolutive, valvidar, &e. (vest. imbricativa, convolutiva, valvaris, &e.), 
but of the leaves or pieces, that they are imbricate, conrolute, valvate, &e., in 
wstivation; but such precision of form will seldom be attended to in botan- 
ical descriptions. 


FIG. 255. Diagrammatic cross section of an unopened tlower of Linden: its outer 
circle of floral leaves (sepals) valvate in the bud; the inner (petals) between convolute 
and imbricate. 


ZESTIVATION, OR PRAEFLORATION. 135 


cover those within. The most familiar case is that of the petals 
of Mignonette and the whole genus Reseda. 

255. Valvate or Valvular, when the margins meet squarely in the 
bud, without any overlapping, like the valves of a dehiscent cap- 
sule. Familiar examples are aiforded by the calyx of the Linden 
(Fig. 255) ; also that of the Mallow, Rhamnus, Fuchsia, and the 
whole of the seyeral natural orders to which these belong. A 
modification of this, caused by some induplication or involution 
of the edges of the individual leaves, occurs in most species of 


Clematis: in Clematis Virginiana, they Gf) 
merely project within (valvate-indupli- (2 ot 
cate); in Clematis Viticella, they are Dad 0 


conspicuously involute (valvate-involute), —— 

or yalvate with margins involute. Some- oa a 
times (as in the calyx of certain Malvacew) the joined edges 
project outwardly (or are valvate with reduplicate margins), but 
only slightly so. 

256. Imbricate or Imbricative is the general name for zstiva- 
tion (or yernation) with overlapping. The name is taken from 
the overlapping of tiles or shingles on a roof, so as to break 
joints or cover edges. It was first applied, by Linnzeus, to 
leaves or scales on a stem, when thickly set and incumbent in suc- 
cessive ranks or heights, the upper partly covered by those next 
below. ‘The involucre of an Aster or of the common Sunflower 
is a typical illustration ; as also the leaves of a Camellia-fiower, 
the sepals as well as the petals; and the sepals or outer leaves 


of a Flax or a Geranium-flower afford a simpler but similar 
instance, although, from the parts being nearly of the same size 
and at the same height, the overlapping is lateral instead of 
obviously from below. Fig. 258, 259, and the outer part of 
260, also the inner leafy circle of 255, illustrate in diagram this 
true and simple imbricative sestivation of a definite number of 


FIG. 256. Valvate-induplicate flower-leaves (calyx) of Clematis Virginiana, &e. 
257. Valvate-involute, asin C. Viticella. 

FIG. 258-260. Imbricate zestivation: 258, in two whorls of three leaves each (calyx 
and corolla); 260, same of five leaves in the outer circle, those of the inner circle con- 
volute; 259, a single set of three imbricated leaves (in the corolla of Magnolia), 
almost completely encircling each other. 


136 PHYLLOTAXY, OR LEAF-ARRANGEMENT. 


parts.!_ It is characteristic of it that some parts (one or more) 
are wholly exterior or covering in the bud, and others (one at 
least) interior or covered, at least the margins. Imbricative 
eestivation, it will be seen, naturally attends alternate or spiral 
phyllotaxy (248, and see Fig. 242, 243); and if it be main- 
tained that these sets of three, five, &c., in the blossom are not 
depressed spirals, but whorls or cycles (as may commonly be 
the case in the corolla, but hardly in the calyx), it is not less 
true that the parts are apt to comport themselves in the exact 
manner of a depressed spiral. The kinds of regular imbrication 
of alternate leaves, &c., may be specified by the terms or frac- 
tions expressive of the particular grade of phyllotaxy (3, 4, 2, 
3, &c.). But some of them have received special names, which 
may be employed, as subordinate to the general denomination of 
imbricate. The most important of these are the 

Equitant, where leaves override, the older successively astride 
the next younger. The typical instance is that of ancipital or 
two-ranked (4) conduplicate leaves, successively clasping, at least 
next the base, as in Iris, Fig. 217. In what Linnzeus termed 
equitant-triquetrous (well seen in Fig. 240, 241), the leaves are 
three-ranked (being of the 4 order), and each imperfectly 
conduplicate. 

Quincuncial sestivation (as in the outer part of Fig. 260) is 
simply the imbricate estivation of five leaves (2), in which 
necessarily the first and second are external, the fourth and fifth 
internal, and the third with one margin external, where it over- 
lies the fifth, and the other internal, where it is covered by 
the first. 

Alternative vestivation, as already stated (252), comes from 
verticillate or cyclic phyllotaxy, and the alternation of successive 
whorls. When two such whorls, say of three leaves each (as in 
Fig. 258), are so condensed or combined as to form apparently 
one set or circle of six members (as in the flower-leaves of most 
Liliaceze), three members alternate with and are covered by the 
other three, and this sort of imbricate sestivation is produced. 
More properly, the two series are to be considered separately. 
Where the parts are four (as in Fig. 395), the normal imbrica- 
tion is decussate, two exterior and two interior. This is some- 


1 All the examples referred to result from alternate or spiral phyllotaxy, 
the former of higher series, the latter of the 4 (Fig. 258, 259), and of the 2 
(Fig. 260) order. Instead of separating (with DeCandolle and others) the 
2 arrangement as different in kind from»the imbricate (under the name of 
quincuncial estivation), we should count it as a typical case. Otherwise the 


¢ arrangement might equally claim a generic distinction, also the 3, &e. 


ZESTIVATION, OR PRA FLORATION. 137 


times a clear case of binary instead of quaternary, 7.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 2 order with one piece omitted, and 
the others adjusted so as to fill the space. 

257. There are various deviations from normally imbricative 
estivation, 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 sestivation, because the individual leaves are involute 
in a manner approaching the convolute vernation of Linnzeus. 
Another is the Vexillar, as in the Pea tribe (Fig. 306), where 
members which should be external have somehow developed as 
internal, both in calyxand corolla. A third (which has received 
the usually quite meaningless name of Cochlear, 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 


@ i) @ 

261 262 263 
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- 
cate.’ To bring Fig. 261 back to the quincuncially imbricate 


1 Tt would not be amiss, therefore, to name one of these modes, viz. that 
of Fig. 261, Subimbricate, and the other, Fig. 262, Subconvolute. George 


FIG. 261. Quincuncial imbricate mo lified 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) «stivation, in diagram. 
Tn these three diagrams, the dark circle above represents the position of the axis, the 
flowers being axillary. 


15 PHYLLOTAXY, OR LEAF-ARRANGEMENT. 


form, we have only to reverse a single overlapping on the left- 
hand side of the figure. To restore Fig. 262 to the convolute, 
we have only to reverse a single overlapping at the lower right- 
hand 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 different flowers on the same 
individual, thus indicating an easy passage between the imbricate 
zestivation in the proper sense and the 

258. Convolufe, otherwise called Obvolute or Contorted, or 
Twisted, Fig. 263, and inner circle of Fig. 260. Here each leaf 
successively overlaps a preceding and is overlapped by a following 
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 are 
broad and much overlapped, as in Fig. 264. Brown 
( included this among the forms of imbricate zestiva- 
( ey, tion, and so does Eichler, particularly distinguishing 
St it, however, under the name here preferred. The 

264 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 Malvacez, 
Onagracez, Apocynacez, Gentianaceze, Polemoniacez, &c.), and 
is so distinct in its nature, that it may well take rank among 
the primary kinds of zstivation. 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 others), the convolute 
and the valvate (having all the members of the series on the 
same plane) answer to verticillate phyllotaxy, or to whorls 
instead of depressed spirals." The name which this mode of 


i 


if 
(: 


Henslow, in Trans. Linn. Soc. ser. 2, i. 178, proposes to call the former half- 
imbricate; the latter (following the faulty French example) is his imbricate 
proper. 

The subimbricate mode has two varieties, distinguished by Eichler (in 
Bliithendiagramme) as ascensive, when the lower or anterior (7. e. the pieces 
next the subtending bract or leaf) are successively exterior (as in Fig. 261), 
and descensive, when the covering is from the upper side, i. e. from the side 
next the axis. 

1 Still, as those members of a quincunx which normally should be wholly 
external do sometimes 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 five overlappings would have 
to be reversed. 


FIG. 264. Convolute (also called contorted) wstivation of a corolla. 


ZESTIVATION, OR PRAEFLORATION. 139 


zestivation ought to bear is not yet well settled, but that of con- 
volute, here preferred, will probably prevail.’ 

259. In recapitulation, these principal forms of the xstivation 
of floral circles may be classified in a synopsis. They are: I. 
those not closed = open or indelerminale: I. closed; and these 


1, with the margins not overlapping = valeate ; 2, with margins 
overlapping ; a, one or more with both margins covered = Bitbrs t- 


cate; 6, all with one margin ‘covered, the other uncovered = 
convolute. 

260. Plicate or Plaited, when applied to the flower-bud as a 
whole, is ina somewhat different category. The term is here 
used for the plaiting 


of a tube or cup, 

composed of a circle 

of leaves ‘eae 
oT 


into one body. 

is well marked in 
the corolla of as alpaind 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 Conyolvulus and Datura (Fig. 266-268), 
the narrow plaits overlap one another in a con- 
volute way, when they are said to be Supercolute. 
In the common Morning Glory and some other 
species of Ipomoea, these plaits are besides spirally twisted or 


1 See article entitled “ Hstivation and its Terminology,” above referred to. 
The earliest name is Obvolute, given by Linneus to the kind of vernation in 
which two leaves (conduplicate ones in his diagram) are put together so 
that one half of each 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 estivation, 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 o) 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 Ipomeea) ; 
that really no twisting accompanies the overlapping in a majority of cases 
of this xstivation; 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 inthe 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 are twisted to the left.’ 

261. 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 estivation. It 
may be either to the right (dextrorse) or to the left (sinistrorse). 
The application of this term depends upon the assumed position 
of the observer, whether outside or inside. We 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 
Fig. 267, 268, dextrorse.? 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 estivation in such cases as deztrorsum 
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 estivation 
(as by Jussieu to the petals of Malvaviscus), but without definition in this 
sense ; it has for many years been steadily adopted by the present 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 Linnzus 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 
Linneus 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 Linnzeus 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 
Peduneles and Pedicels, in the latter to 
Bracts and Lractlets. 

264. A Braet (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.’ 
They are commonly, but not always, 
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 269 
minute and functionless, sometimes obsolete (as in Cruciferae) , 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 (spathe) of Indian Turnip, partly cut away below to show the 
fleshy spike (spadix) of flowers wlich it surrounds. 


142 ANTHOTAXY, OR INFLORESCENCE. 


fugacious. Each flower is subtended by (grows from the axil 
of) a bract in Fig. 277-280, &e. A cluster of flowers is sub- 
tended by a conspicuous and colored bract in Fig. 269, 270, 271 ; 
by a circle of colored bracts, imitating white petals, in Fig. 294. 

Spatne is the name given to such an enclosing bract, or 
to two or more leaves successively enclosing a flower-cluster. 


Ixvo.ucre 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 Hepatica and 
Mallow. A compound inflorescence may have both a genera 
and a partial involucre, one for the general flower-cluster, otheis 
subtending the partial clusters. The name of involucre is then 
reserved for the general one; that of 

INVOLUCEL is applied to the partial, secondary, or ultimate 
involucres. 

Bractiets (Lat. Bracteola, diminutive of bract) are bracts of 
e secondary or ultimate order. For example. in the slender 
flower-cluster, Fig. 277, 6 is a bract, subtending each individual 
flower-stalk ; b’ is a bractlet, or bract of secondary order, borne 
on that partial flower-stalk itself. The French naturally translate 
the Latin bracteola into bractéole (pl. bractéoles): in English, 
bractlet is an idiomatic and better diminutive. 

Patets (Lat. Palee), also called Chaff, are diminutive or 


FIG. 270. Monophyllous spathe of Indian Turnip, with tip more erect. 271. Spathe 
and spadix of Calla. 272. Raceme of Cherry, leafy at base. 273. Dichotomous cyme. 
274. Panicle of Meadow-Grass, 275. A corymb. 


BRACTS AND FLOWER-STALKS. 143 


chaff-like bracts or bractlets on the axis (or receptacle) and 
among the flowers of a dense inflorescence, such as a head of 
Composite (275, Fig. 287, 288); and the name is also given 
to an inner series of the 

Guiumes of Grasses. These are peculiar chaffy bracts or bract- 
lets which characterize the inflorescence of Grasses and Sedges. 

265. Peduncle is the general name of a flower-stalk, that is, 
of an axis or stem, which instead of foliage, or at least ordi-_ 
nary foliage, supports a 
flower-cluster or a single 
flower. In Fig. 276, 
each peduncle (rising 
from the axil of an ordi- 
nary leaf, and therefore 
answering to a branch) 
bears a solitary flower. In Fig. 277, the peduncle bears a series 
of flowers, or a flower-cluster... In this instance, each flower is 
borne on a flower-stalk of its own, that is, upon a 

PepiceL. This is the name given to distinguish 
a partial flower-stalk, or, more strictly, the stalk 
of each individual flower of an inflorescence. (Fig. 
277-284.) In less simple flower-clusters, with 
ramification of two, three, or more grades, general 
peduncle, partial peduncles, and pedicels have to be 
distinguished: the term pedicel is reserved for the 
ultimate ramification. 

Scape is the name given to a peduncle rising 
from the ground, as that of most Primulas, of 
Dodecatheon, Hepatica, and the so-called acaules- 
cent or stemless Violets. 

Ruacuis (backbone) is a name given to the axis 
of inflorescence; that is, the continuation of the 
stem or peduncle through a somewhat elongated 
flower-cluster, as in a spike of Birch or of Plan- 
tain, Fig. 289,290. When this axis is short, as in 
a head (Fig. 285-288), it is usually called the 
RECEPTACLE, a word also used for the axis or cauline- 277 
part of a flower. The context should show when receptacle of 
inflorescence, and when receptacle of the flower itself, is meant. 
Both belong to axis or stem. 


reo 

¢ 
ae) 
e 


FIG. 276. Moneywort, Lysimachia nummularia, with axillary one-flowered 
peduncles. 

FIG. 277. A Raceme, with a general ee (p), pedicels (p’), bracts (b), and 
bractlets (0). 


144 ANTHOTAXY, OR INFLORESCENCE. ° 


266. Position of Flowers or Clusters. Flower-buds accord 
with leaf-buds in origin, position, and structure, to this extent at 
least, that the parts of both are leaves or homologues of leaves, 
crowded in whorls or spirals upon a short portion of stem or 
axis; aud as leaf-buds are either terminal or axillary (15, 75) 
so also are flower-buds ; as a leaf-bud may give rise to a simple 
or a compound growth, 7. e. may branch again and again, or not 
branch at all, so flower-bearing branches, or the flower-bearing 
extremity of a stem or branch, may bear a single flower, or a 
more or less compound cluster. Thus, in Fig. 276, an axillary 
peduncle, or naked branch, bears at apex a solitary flower; in 
Tig. 277, a peduncle bears a loose cluster of flowers, each of 
which springs from the axil of a small bract; in Fig. 285, 
a terminal peduncle bears at summit a dense flower-cluster. 
Flowers are either solitary or in clusters. When solitary, they 
are naturally without bracts, being subtended instead (as in Fig. 
276) by ordinary foliage. 

267. ‘The elevation either of a solitary flower or a cluster on a 
peduncle, or of individual flowers of a cluster on pedicels, is only 
incidental. ‘The flowers may be stalkless, i. e. sessile. 

268. The Kinds of Inflorescence which have received distinctive 
names are various, but are all reducible to two types, which, 
generally well marked, may sometimes pass into each other, and 
which are not rarely combined in the same compound inflores- 
cence.’ The two types differ in basis as do axillary from ter- 
minal buds ; in the one the flowers are axillary or lateral, in the 
other terminal in respect to the axis from which each flower or 
its pedicel arises. But inasmuch as every flower, whatever its 
position, is terminal to its own stalk or axis, it is better to dis- 
tinguish the two types in other terms, and to name them the 

269. Indefinite and Definite, or, in equivalent and similar terms, 
the Indeterminate and Determinate.2 Each may be either simple 
or compound. It is from the simple that the definitions are to 
be drawn. In the former type, the rhachis or main axis of the 
inflorescence is not terminated by a flower, but lateral axes, or 
pedicels, are. In the latter, both the main or primary and the 
lateral or secondary axes or stalks are so terminated. An inde- 
terminate flower-cluster may go on to develop internode after 


1 Inflorescence, as has been well insisted on by Guillaud (in Bull. Soe. 
Bot. France, iv. 29), is a mode, not a thing. The things sometimes but in- 
appropriately so called are flower-clusters, for which, if a general technical 
name is needed, that of Anthemia, in English Anithemy, suggested by Guillaud, 
is as good as any. 

2 Also named by Eichler (Bliithendiagramme, 33, following Guillaud, 1. c.) 
the Cymose and the Botryose type. 


THE TWO TYPES. 145 


internode of axis, and one or more leaves (bracts) at each node, 
and then a flower in the axil of each bract, until its strength or 
capability is exhausted. Or it may stop short with very few 
flowers ; but the uppermost and youngest one will not really ter- 


270 289 

minate the rhachis (7. e. come from a terminal bud), though i: 
may appear to do so. (Fig. 272, 277-279, &e.) The lower 
flower-buds are evidently the oldest, and accordingly the first to 
expand; and the expansion will proceed regularly from below 
upward: wherefore this type of inflorescence has been called the 
Ascending ov Acropeta!l ; likewise the Centripetal, because, when 
the flowers are brought to the same level or near it (as in Fig. 
279, 280) by a lengthening of the lower pedicels, with or with- 
out relative shortening of the rhachis, the evolution 
is seen to proceed from circumference to centre. 
There is thus no lack of names; but, inasmuch as 
the following type is commonly referred to under the 
general name of Cymose, to this has recently been 
given the counterpart name of Botryose. (271.) 


281 


270. A determinate flower-cluster (as seen in its gradual 
development which is not rarely presented) has the last internode 


FIG. 278-280. Diagrams of indefinite, indeterminate, contripetal, or botryose in- 
florescence: 278, Raceme; 279, Corymb; 280, Umbel. 

FIG. 281-284. Diagrams of definite, determinate, centrifugal, or cymose inflores- 
cence: 281, a false or descending raceme; 282, a solitary terminal flower; 283, same with 
two lateral flowers developing, forming a 3-flowered cyme; 284, same with lateral 
peduncles 3-flowered, or a pair of 3-flowered cymes, beside the central or primary 
terminal flower. 


146 ANTHOTAXY, OR INFLORESCENCE. 


of its axis terminated by a flower (Fig. 281-284), which answers 
to a terminal bud. If more flowers appear, so as 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 
by the size of the flower-buds or degree of expansion of the 
blossoms. Fig. 281 best shows why a determinate or definite in- 
florescence is sometimes said to be Descending ; Fig. 283 shows 
why it is called Centrifugal, 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 Linnzeus, 
but defined without reference to the order of evolution of the 
flowers. They are the Raceme, Corymb, and Umbel, with flowers 
raised on pedicels; the Spike and Head, with sessile flowers ; 
also some modifications of these, notably the Ament and the 
Spadix. The raceme may be taken as the type. Sotrys is 
equivalent to racemus, &c.; and, as the type includes diversity 
of forms to which the name racemose would seem inapplicable, 
the term botryose (botrytischen of Eichler) is best 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. 

273. 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 be- 
coming flat-topped or convex. ‘The centripetal character is thus 
made apparent. The greater number of the corymbs of Linnzeus 
and succeeding botanists are cymes, the central flower first ex- 
panding. And the term corymbose or corymb-like is still much 
used in descriptive botany for a ramification which is mainly of 
the cymose type, and where in strictness the term cymose should 
be employed. 

274. An Umbel (Fig. 280), as in Asclepias, &c., differs from 
a corymb only 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 BOTRYOSE 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 Faseiele (or the pedicels 
said to be fuscicled) ; 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 
ahead 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 ons 
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), 7.e. naked of bracts: when 
they are present, it is paleate or chaffy. A peculiar sort of head, 
not undeserving a special name (though this is not necessary 
in descriptive botany), is the 

Antnopium, the so-called Compound Flower of the earlier 
botanists, which gives the name to the vast order of Composite. 


FIG. 285. Cephalanthus occidentalis, the Button-bush; a pair of leaves, and a 
terminal peduncle bearing a dense head (capitulum) 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 (/igulate) corollas of the several flowers imitating 
the petals of a single blossom. In some (such as Dandelion 
and the Cichory, Fig. 286), all 
the flowers of the head bear 
these petal-like corollas; in 
more (such as Aster, Sun- 
flower, and Coreopsis, Fig. 
287), only an outer circle of 
flowers does so; the remain- 
der, smaller and filling the 
centre (or disk), 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 depressed, 
bearing the flowers on what then becomes the upper surface, 
which adds to the imitation.? 

Syconium. 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 


1 The receptacle of an Anthodium has been termed Clinanthium or Phor- 
anthium ; and its involucre, a Periphoranthium or Periclinium. 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 morpho.ogy is ex- 
plained and illustrated. Viewed as an inflorescence, it has also 


been named a HypantTHopium. 


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. : — 

Spapix, 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 


290 


a spathe, and there is nospadix. 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 (Ul); also one ligulate 
and neutral ray-flower and part of another (c,¢): in d, d, the bracts or leaves of the 


involucre are seen in section. 


FIG. 289. Catkin of White Pirch. 290. Young spike of Plantago major. 


150 ANTHOTAXY, OR INFLORESCENCE. 


Ament or Catkry. This is merely that kind of spike with 
scaly bracts borne by 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. 

277. Any of these forms of simple inflorescence may be com- 
pounded. Racemes may themselves be disposed in racemes, spikes 
in spikes (as in Triticum), heads 
be aggregated in heads, umbels in 
umbels, corymbs may be corym- 
bosely compound, &e.; forming 
compound racemcs, spikes, umbels, 
and the like, the terminology of 
which is easy. The most usual 
case of truly homomorphous com- 
pounding is that of umbels ; the 
inflorescence of much the larger part of Umbelliferae being in 
compound umbels, as in Fig. 290%. There is then the general 
umbel, the rays of which become peduncles to 
tue partial umbels, and the rays of the latter 
are pedicels. Umbella and Umbellule desig- 
nate in Latin terminology the general and its 
partial umbels. Umbellets (coined by the late 
Dr. Darlington) may well replace the latter 
as the English diminutive. But umbels are 
sometimes racemosely arranged, as in Aralia 
spinosa, heads may be arranged in spikes, 
and so on. 

278. A Panicle, of the simple and normal 
sort (as illustrated in Fig. 291), is produced 
when a raceme becomes irregularly compound 
by some (usually the lower) of its pedicels 
developing into peduncles carrying several 
flowers, or more than one, or branching again 
and again in the same order. But in com- 
pound clusters generally the secondary and 
tertiary ramifications are apt to differ in type 
as well as in particular mode, giving rise to 

291 heteromorphous or mixed inflorescence. (288.) 
As Linneeus defined the term, and as it has generally been em- 
ployed in botanical descriptions, the panicle is a general term 


FIG. 290¢ Compound umbel of Caraway. 291. A simple panicle. 


THE CYMOSE TYPE. aaa 


forvany loose and diversely branched cluster, with pedicellate 
flowers. It is therefore difficult to restrict it in practice to the 
indeterminate type. . 

279. Varieties of Determinate or Cymose Inflorescence. The 
plan of this type has been sufficiently explained. (270.) Its 
simplest condition is that of a solitary terminal flower, peduncu- 
late or pedicellate (as in Fig. 282), or sessile. The production 
of more flowers, necessitates new axes from beneath, from the 
axils of adjacent leaves or bracts. ‘These, being later, render 
the evolution centrifugal. The simplest flower-cluster (unless 
we call the solitary flower of Fig. 282 a one-flowered cluster) is 
that of Fig. 283, where a secondary floral axis or peduncle has 
developed from the axil of each leaf of the uppermost pair, or 
where with alternate leaves there is a single uppermost leaf, and 
then only one such peduncle, and thus is produced a three- (or 
two-) flowered cymose cluster. The flower of the primary axis 
is marked by its bractless peduncle (therefore a pedicel) ; the 
lateral and secondary peduncles are known (commonly or nor- 
mally) by their bracts or bract; the portion below the bracts 
is proper peduncle; that above, of single internode, pedicel. 
Bracts, like other leaves, have potential bud§ in their axils ; these 
in an inflorescence give the third order of ramification, each 
branch tipped with its flower; and so on. 

280. The Cyme is the general name of this kind of flower-cluster 
in its various forms. One of these very simple cymes, by itself or 
as a part of a larger cyme, may be called a Cymule. The regular 


292 


cyme usually accompanies opposite or other grades of verticillate 
leaves, but is not rare in the alternate arrangement. It is 
readiest understood in an opposite-leaved plant with regular 
opposite ramification, as in an Arenaria, Fig. 292. By its con- 
stitution, a cyme proceeds from simple to compound. It mat- 


FIG. 292. Dichotomous or biparous cyme (cyme bipare of Bravais, Dichasium of 
Eichler) of Arenaria Michauxii. 


152 ANTHOTAXY, OR INFLORESCENCE. 


ters little whether its development is progressive, the flowers of 
the ultimate ramifications expanding after the earlier have matured 
fruit, and with subtending bracts conspicuous or foliaceous ; or 
whether, as in Elder and Hydrangea (Fig. 293, and in Fig. 
273), the bracts are minute and caducous or abortive, and the 
ramification complete with all the flower-buds well formed before 
the oldest expand, so that the whole is in blossom almost at the 
same time. But a cyme may be properly said to be compound 
when the primary axis in it is a peduncle instead of a pedicel, 


293 


and supports a cluster (cyme or cymule) instead of a solitary 
central flower at the main divisions.t One form of the regular 
cyme, on account of its compactness, is named the 

GromeruLe. This is merely a cymose inflorescence, of any 
sort, which is condensed into the form of a head, or approach- 
ing it. Of this kind is the so-called head of Cornus florida, and 
of the herbaceous C. Canadensis (Fig. 294), which shows the 


1 The dichotomous or two-branched cyme is the commonest, but is some- 
times marked by suppression of internodes; as, for example, where the 
branches are apparently in fours, in an umbelliform way ; but these are two 
sets of two, with the internode between the pairs extremely short; or where, 
as in Elder, the branches or rays are five, in this case consisting of the same 
two pairs and a central one, which is a many-flowered continuation of the 
primary axis. Or 5-rayed cymes, &c., may be founded upon alternate leaves 
with shortened internodes, the rays or peduncles axillary to them thus 
brought into an apparent whorl. 

Bravais distinguished cymes as mu/tiparos, with three or more lateral 
axes ; biparous, with two, and uniparous, with only one (cyme multipare, 
bipare, unipare). To these Eichler gives the substantive names, severally, 
of Pleiochasium, Dichasium, and Monochasinm. Only the latter needs illustra- 
tion ; the others being as it were compounds of this. 


FIG. 293. Compound cyme of Hydrangea; with some neutral and enlarged mar- 


ginal flowers. 
“ 


THE CYMOSE TYPE. 158 


composition best, on close examination. A condensed but less 
capitate cyme, or cluster of cymes, was called by Raper and 
DeCandolle a FascrcLe; and this terminology has been much 
adopted. It is properly enough 
said to be a fascicle, which, as 
used by Linneus and others, 
means a bundle, or close collection 
of parts, whether leaves, pedun- 
cles, or flowers; but a fascicte is 
not necessarily a cyme (274), noris 
there need of a special substantive 
name for a compact cyme, which 
may either be simply so called or 
it may pass into the glomerule. 
281. Botryoidal forms of Cymose 
Type, or False Racemes, &c. The 
regular cyme seldom continues 
with all its ramifications. In 
Fig. 292, after the second forking, 
one of the two lateral peduncles 
mostly fails to appear, and in some } a4 i 
parts one of the bracts also; and ultimately the lateral peduncle 
present is bractless, like the central, therefore equally incapable 
of further ramification, being reduced to a pedicel of a single inter- 
node. This suppression some- 
times begins at the first fork- 
ing or at the very base; and, 
when followed throughout, it 
reduces a biparous or dichoto- 
mous cyme to one half, and, 
converts this half (when the 
axis straightens) into the sem- 
blance of a raceme if the 
flowers are pedicelled, or of a 
spike when they are sessile. 296 27 
Fig. 296 is a diagram of such an inflorescence as that of Fig. 292. 
with one lateral branch uniformly suppressed at each division, the 
wanting members indicated by short dotted lines. Cases exem- 
plifying this occur in portions of the inflorescence of some of our 


FIG. 294. Plant of Cornus Canadensis: flowering stem bearing a cluster of leaves 
aboye, then continued into a peduncle, and terminated by a glomerule of very small 
flowers; this subtended by a colored and corolla-like involucre of four bracts. 295. One 
of the flowers taken from the glomerule, enlarged. 

FIG. 296. Uniparous cyme or sympodial false raceme, with opposite leaves or bracts. 

FIG, 297. Form of the same, with alternate leaves or bracts. 


154 ANTHOTAXY, OR INFLORESCENCE. 


smaller Hypericums, and notably in H. Sarothra, in which the 
leaves are all reduced to bracts. It is not always easy to show 
why this is nota 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, which commonly runs into a coil, finds explanation in the view 
that each portion is the lateral branch from the axil of the subtend- 
ing leaf: and occasionally the other axil produces a similar one, 
thus revealing the cymose 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. 296, 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. Sympodial forms. The explanation is that the axis of 
inflorescence in such cases, continuous as it appears to be, is 
not a simple one, is not a monopode, but a synypode (110, 116, 
notes), z.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. 297), 
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 strictly to one 
side and the flowers to the other, in Fig. 297, shows that these 
leaves 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 Houseleek, &c. 

283. A further difficulty in the morphology of clusters of this 
class comes from the early abortion or complete suppression of 
bracts. This is not unknown in botryose inflorescence, occurring 
in the racemes, of almost all Cruciferee : it is very common in the 
cymose of all varieties, and especially in the uniparous ones in 
question, which characterize or abound in Borraginacexze, Hydro- 
phyllaceze, 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, &e. 

284. Commonly these false racemes or spikes (or botryoidal 
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 scorpzord. 
As the coil is a helix, it has also been named helicoid.1 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.? 


1 Scorpioid and Helicoid have been carefully distinguished by later 
morphologists, 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: PLe1ocnasium, the multiparous cyme ot 
Bravais. 

B. Lateral axes two: Dicnasium, the Liparous cyme of Bravais. 

y. Lateral axis one: Monocuasium, the uniparous cyme of Bravais. 


The latter, or the corresponding divisions of the preceding sorts, may be 
divided as follows: 


156 ANTHOTAXY, OR INFLORESCENCE. 


286. Sundry complications and obscurities are occasionally 
encountered in anthotaxy or phyllotaxy, which cannot here be 


x 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: ScuravuBet [screwlike] or Bostryx 
[ringlet or curl], the uniparous helicoid cyme of Bravais. 

2. Lateral axes falling alternately on opposite sides of the relatively main 
axis: WicKEL or CINcINNUS [a curl], the uniparous scorpioid cyme of 
Brayais. 

* * Lateral axes medial [in the same plane] relative to the main axis. 

3. Lateral axes in successive generations always on the back side of the 
axis from which it springs: FXcneL, Ruiprpium [fan]. 

4. Lateral axes in successive generations always on the upper side of the 
axis from which it springs: S1cneL, Drepanivm [sickle]. 

The subjoined simple diagrams from Eichler (Fig. 298-503) illustrate 
these forms. The ramification is, given without the bracts, which theoreti- 
cally or actually subtend the axes of each generation. The student may add 
them, and so more readi:y apprehend the characters. 

Eichler recognizes the forms with median (antero-posterior) position of 
axes in Monocotyledons only. 
It is natural to distichous phyl- 
lotaxy, and it accords with 
the general rule that, in mono- 
cotyledonous plants, the first 
leaf of the branch, or the com- 
monly solitary bractlet of the 
° peduncle, stands over against 
70 and facing the bract or leaf 
28 from the axil of which said 
branch or peduncle springs, 7.e. is posterior and next the parent axis, as 
shown in the diagram, Fig. 304, 305. 


301 300 a be 
: \ 
OQ-O-0-0-0~ OKs pp 
ps = 0 ( 0) 
a Oana 


FIG. 298. Diagram of the Cincinnus. 299. Diagram of the Bostryx The flower- 
axes numbered in succession. 

FIG. 300. Diagram of the Rhipidium. 302. Ground plan, indicating the order of 
evolution of the flowers. 

FIG. 301. Diagram of 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; is bract, and b/ bractlet. 


THE CYMOSE TYPE. VT 


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 
Fichler’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 Borraginacee and 
Hydrophyllacexw, &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, &«., 
and its flowers are commonly one-ranked. In the Cincinnus or true scorpioid 
_ eyme (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 (b’) of each successive secondary axis (a’) stands next the 
axis (a) and over against the bract (4) 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 ‘Hy dro- 
phyllaceous species, in Houseleek, Tradescantia erecta, &c. This comes 
through antidromy, that is, the phyllotaxy of aC) 
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 T 
right alternately. Fig. 301 is a plan of this Ls 
two-ranked unilateral arrangement. Whennot 


too crowded, both Cincinnus and Rhipidium ‘ : 

are apt to have a zigzag rhachis. ( IT ao. 
These two last-mentioned kinds are so gen- a vie FA 

erally alike in character, as are equally the ys 

Bostryx and the Drepanium, that the four spe- i 

cies may as well be reduced to two. As these 7. Ge ) = 

severally include the scorpioid and the helicoid a ~ Vy, - 

uniparous cymes of modern anthotaxy, these v Vo) 

terms may be retained to designate them. Or, 

if other terms in use be preferred to scorpioid Pd athe 

and helicoid, the form with two-ranked flowers B 

may be denominated Cincinnal, that with single- 2057 


ranked Gostrychoidal. 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 are all 
absent, no obvious external difference remains. 


FIG. 305%. Ground plan (from Eichler’s Bliithendiagramme, i. 38) of the scorpioid 
inflorescence of Tradescantia erecta, between bract (B) below and axis (a) above: 
I., IL., IIL, &c., the successive flowers: v' is the bractlet of the first and bract of the 
second flower, and so the others in succession up to 7° 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.’ 

287. Mixed Inflorescence is not uncommon. This name is 
given to clusters or ramifications in which the two types are con- 
joined. Being heteromorphous, they are almost necessarily com- 
pound, the two types belonging to different orders of ramification. 
But under it may be included cases of comparatively simple 
inflorescence, at least in the beginning, some of which nearly 
fuse the two types intoone. 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 wovld answer to Fig. 281, which, 
blossoming from above downward by simple uniflorous lateral 
axes along a monopodial primary axis, is a simple racemiform 
eyme, 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 
the 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 spicata 
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. 281 the lower pedicels were prolonged 
to the level of the upper, a simple corymbiform cyme would be 
seen, with simple centrifugal evolution, that is, regularly from 
the centre to the margin; thisis 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 the side of a bract in false 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 umbel of two 
flowers, transverse to the bract (as in many species of Desmodium), and 
thus seemingly lateral to it, or to a single flower 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 (concaulescence of Schimper), is common. So likewise bracts 
or leaves may be for a good distance adnate to sympodial shoots, whether 
peduncles or leafy flowerless branches. This (named _ recaulescence by 
Schimper) is of most frequent occurrence in Solanacee (in Datura, Atropa, 
most species of Solanum, &c.), and is tie explanation of their so-called 
geminate leaves,-where a large leaf (reaily belonging lower down) has a 
small leaf by the side of it. See Wydler in Bot. Zeit. ii. 689, &c., Sendtner 
in Fl. Bras. x. 183, and Eichler, Bliithend. 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 generat 
and of the partial anthotaxy, or that of the main axis of inflo- 
rescence and that of its ensuing ramification.' 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 Labiatsze and most Scrophulariaceze, 
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 

Tuyrsus. 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 

Mrxep Panicre. 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. So 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 Labiate. 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 Cymo-otryes the mixed inflorescence 
composed of cymes developed in botryose order, 7.e. the thyrsus; and 
Botry-Cymes, the reverse case of racemes, &c., cymoscly aggregated. For 
the former, the old name thyrsus serves appropriately and well. 


160 ANTHOTAXY, GIR} 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. (515.) 

290. Anterior and Posterior, otherwise called /nferior and 
Superior, and therefore Lower and Upper,’ are primary relations 
of position of an axillary flower with respect to subtending bract 
and the axis to which the bract pertains. The flower is placed 
between the two. The portion of the flower which faces the 
subtending bract is the anterior, likewise called inferior or lower. 

The opposite portion which faces the axis of 

5 inflorescence is the posterior, or superior, or 

upper. ‘The right and left sides are lateral. 

~N (Fig. 304, 306.) These relations do not 

v( f =) ee in a solitary flower terminating a 
g simple stem; but when such an axis produces 

©) axillary branches with a terminal flower, the 

———4 relation of this flower to the preceding axis 

Lm & and its leaf is manifest, just as in indetermi- 
nate inflorescence. 

291. Median and Transverse. The position of parts which lie 
in antero-posterior line, or between bract and axis, is median. 
Thus, in Fig. 504 and 305, the parts are all in the median plane : 
in Fig. 306, the btactlets, b’, b’, are lateral or collateral, or (being 
in the opposite plane) transverse. 

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. 304, 305) ; 
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.2 Commonly there is only this 


1 Not (with propriety, although the terms have been so used) exterior or 
outer for the anterior, and interior or inner for the posterior position. These 
terms should be 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 Bracteole: 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 case 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 bractlet 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. Diagrain (cross section) of papilionaceous flower and its relation to axis 
(a), bract (b), and braetlets (b7, 67). 


THE CYMOSE TYPE. 161 


posterior one to a simple axis in Monocotyledons, and two 
transverse ones in Dicotyledons, 7.e. one to the right and the 
other to the left of the subtending bract, Fig. 306, 4’ b’.. When 
the latter form a pair, they are perhaps always truly transverse ; 
when alternate, they stand more or less on the opposite sides and 
transverse. When more than one in Monocotyledons, they may 
become either median or transverse, or even intermediate. The 
relation of bractlets 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 ;! 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 bractiets, right and left, the first 
flower-leaf is at } divergence from one (the uppermost) of them 
when the circle is of three, or at 2 when of five members, or near 
it; but with many exceptions.” 


A tabular view of the kinds of inflorescence and their termi- 
nology, serving as a key, may aid the student.® 


latter, being the leaves which the new axis first bears, Vorb/éftter, 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. 

2 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 M1xep, of the two types 
combined. The Types are: 
I. Main axis not arrested and terminated by a flower. Indeterminate, Indefi- 
nite, Acropetal or Ascending, Centripetal, or BorRYOSE. 
II. Main and lateral axes arrested and terminated by a flower. Determinate, 
Definite, Descending, Centrifugal, or Cymosr. 


{62 ANTHOTAXY, OR INFLORESCENCE 


I. BOTRYOSE TYPE. 


1. Simple, with lateral axes unbranched and terminated by a single flower, and 
Flowers on pedicels, 
Of somewhat equal length on a comparatively elongated axis, RACEME. 
The lower ones longer than the upper, and main axis short, . CoryMB. 
Of nearly equal length on an undeveloped main axis, . . UMBEL. 


Flowers sessile'‘on a very Short main axis, << . .. . . » BAD. 
Flowers sessile on a comparatively elongated main axis,. . . SPIKE. 
Acfleshy.spilke‘or head-is/ar Grice s34,. 1c.) 21a) ba) eS Ee 
A sealy-bracted spike isan 5 - = 2 1 > a. = of = ae) SReANIIONIDIOR 
CATKIN. 


Compound, with lateral axes branched once or more, bearing clusters instead of 
single flowers. 
Irregularly racemosely or corymbosely compound,. . PANICLE. 
Homogeneously and regularly compound, as 


Racemes inaraceme, ... .. .. .. . ». ComMPpouND RACEME. 


Corymbs: conymbose,..4). 2.) a) 5) Sh) . )  eComrounpiGonwane 
Umbels inanumbel, . - .... 4.) .. « «5. 4. ComPouxDiUimern 
Spikes spicate, . . 2. = «4c =. «+ » ~ COMPOUND) Spas 


Homogeneously compound, the secondary ramification 
unlike the primary, as Heads racemose, Umbels spiked, 
Spikes panicled, &c. 


If. CYMOSE TYPE. 


1. Simple, with terminal axis of each generation one-flowered. 
Monopodial, the axis of each oer as evidently re- 
solved into branches, . . . . : 7 ew LERUE Onin 
These more thantwo, . . . . iPicivehnetan or MuLTIPAROUS CYME. 
These only two, . . .Dichasium, Dichotomous or BrrArous CYME. 
Sympodial, the apparently simple axis continued by 
a succession of new axes standing end to end, 
Monochasium, False Raceme or Spike, Botryose or UNtpAROuUS CYME. 
Flowers one-ranked on one side of rhachis, HELIcom Untrparous CyME. 
Flowers two-ranked on one side of rhachis, Scorpiouy Unrparous CyME. 


2. Compound, with terminal axes for one or more earlier 
generations bearing a cyme instead of a single 
flower, . .- . .- -. . . - Warious sorts of CompouND CYME. 


Ill. MIXED INFLORESCENCE. 


1. Anomalous simple, with unbranched one-flowered lateral 
axes (287), suchas . . . PARTLY KEVERSED SPIKES OR RACEMES. 
2. Compound, of various combirations, of which there are 
names for the subjoined : — 
Primary inflorescence botryose, with axis elongating; 


secondary cymose, . . 5 ~ « « RYRSUR: 
Pair of such opposite cymes yee caatuent round 
thesmMaincaiShs phere een ey ws - « «© « « WERTICILLASTER: 


Panicle with some of the ramifications ey mose, . . . MIXED PANICLE. 


THE FLOWER. 163 


CHAPTER VI. 
THE FLOWER. 


Section I. Irs Nature, Parts, AND METAMORPHY. 


293. FLower-Bups are homologous with (morphologically an- 
swering to) leaf-buds, and they occupy the same positions. (266.) 
A FrLower is a simple axis or a terminal portion of one, ina 
phznogamous 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, they 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-Linnean 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 Catyx and CoroLia; and the 
Essential Reproductive Organs, which co-operate in the production 
of seed, the STaMENs and PISsTILs. 

296. Floral Envelopes, Perianth, or Perigone, the fioral leaves 
or coverings. The former is a proper English designation of 
these parts, taken collectively. But in descriptive botany, 
where a single word is preferable, sometimes the name perianth 
(Lat. perianthium), sometimes that of per/gone (or perigonium) , 
is used. Pertanthium,'! a Linnzean 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. Pertgonium, 
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, 
as in the petaloideous 
Monocotyledons. Itis 
also used where the 
morphology is ambigu- 
ous. Generally, the floral envelopes are treated distinctively as 
calyx and corolla, one or the other of which (mostly the corolla) 
may be wanting. 

297. The Calyx is the outer set of floral envelopes. That is 
its only definition. Commonly it is more herbaceous or foliaceous 
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 Linneus (and about the same time Ludwig) used it in the sense of a 
proper calyx, yet with some vagueness. Mirbel and Brown established it 
in the sense of the collective floral covering. DeCandolle revived Ehrhart’s 


FIG. 307. The complete flower of a Crassula. 308. Diagram of its cross-section in 
the bud, showing the relative position of its parts The five pieces of the exterior 
circle are sections of the sepals; the next, of the petals; the third, of the stamens 
through their anthers; the innermost, of the five pistils. 

FIG. 309 <A sepal; 310, a petal; 311, a stamen; and 312, a pistil from the flower 
represented in Fig. 307. 


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. 

298. The Corolla is the inner set of floral envelopes, usually (but 
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 Perars.! 

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 ANDRactuM ; the pistils, the Gynasctum.? 

301. The Stamens* are the male or fertilizing 
organs of a flower. A complete stamen (Fig. 311, 
313) consists of Firament (/), the stalk or support, 
and ANTHER (a), a double sac or body of two cells, 313 
side by side, filled with a powdery substance, Potten, which is 
at length discharged, usually through a slit or cleft of each cell. 


well-formed name of perigonium, 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 inner circles, 
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.” 

2 The male household and the female household respectively, terms in- 
~ troduced by Reeper (Linnea, i. 437), in the form of andraceum and gyneeceum ; 
but the diphthong in the latter should also be @ The orthography 
andrecium and gynecium (early adopted by Bentham, in Labiatarum Gen. et 
Spec.) is conformable to the Linnean Monacia, Diecia, &c. 

3 The name (from the Greek and Latin name of the warp of the ancient 
upright loom, and thence used in the sense of threads) was applied, down 
to Tournefort and later, to the filaments; and the anthers were termed 


FIG. 313. Stamen, composed of /, filament, and a, anther, with cells opening 
laterally and discharging pollen. 


166 THE FLOWER. 


302. The Pistils, one or more to the flower, are the female or 
seed-bearing organs.’ A complete pistil is distinguished into 
.e three parts: the Ovary (lati 

Ovarium, Fig. 314, a, shown in verti- 
seeete » cal section, and Fig. 315, by Linnzeus 
named Germen), the hollow portion 
at the base which contains the 


come seeds ; the SryLe (4), or colum- 
nar prolongation of the apex of the 
ovary; and the Stigma (¢), 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 filament of a 
stamen) may be altogether wanting. 


apices. It came in time to be used as now for the whole organ; but Lud- 
wig (Inst. Reg. Veg.), in 1742, apparently first so defined it, and introduced 
the term Anther for the Apex of Ray, or Theca of Grew. 

1 Following Linnzus, this term is here freely used in the plural, and for 
each actual separate member of the gynecium, each organ which has an 
ovary, stigma, and commonly a style. Tournefort, who appears to have 
introduced the word, employed it in the sense of gynecium. Many authors 
define it thus, and then practically eliminate from botany this, one of the 
oldest of its terms, and one by no means superfluous. The typical pistillum 
of Tournefort is that of the Crown Imperial (Inst. i. 69, & tab. 1) and the 
name is from the likeness to a pestle ina mortar. As it soon became im- 
possible to apply the same name to the pistil of a Fritillaria or of a Plum, 
the cluster of such organs in Caltha, and the capitate cluster and receptacle 
of such organs in a Ranunculus or Anemone, Linnzus, and Ludwig before 
him, took the idea of Tournefort’s name, and used it accordingly. 

“ Pistillum est pars interior et media floris, que ex ovario et stylo com- 
ponitur. . . . Ovarium est pars pistilli inferior, que futuri fructus delinea- 
tionem sistit. . . . Stylus est pars pistilli ex ovario centro producta. .. . 
Summitas styli vel ejus partium Stigma dicitur.” Ludwig, Inst. Reg. Veg. 
41-45, 1742. Without mentioning the plural, the pistil is thus defined in a 
way which necessitates its use. Linnzus (in Phil. Bot.) first defines Stamen 
and Pistillum in the singular number, enumerating the three parts of the 
latter, and afterwards (p. 57) declares that “ Pistilla differunt 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, 6, 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. 


OvuLes, or bodies destined to be-- 


ITS NATURE AND PARTS. 167 


303. The Torus, or Receptacle of the flower, also named 
Tuavamts,’ is the axis which bears all the other parts, that 
upon which they are all dc 
(mnediately or immediately) 3 
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 leayes on 
the stem; the calyx from 
the very base, the petals 
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-buds 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- 
blanee, but the ripe pistils open down the inner angle and 
flatten out into a leaf-like form. ‘The adopted theory supposes 
that stamens and pistils, as well as sepals and petals, are homolo- 


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 méme dans toutes les espéces,” &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 Théorie Elémentaire, third edition, writes, 
“Chaque carpel est un petit tout, un pistil entier, composé d’un ovaire, @un 
style, et dun stigmate.” Of English authors, no other need be cited than 
Robert Brown. The terms in question, then, are : — 

Gynecium, the female system of a flower, taken as a whole. 

Pistil, each separate member of the gynecium; 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 Carpophyll, 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, Linnzus. 
Thorus, 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; 6, petal; c, stamen; 4, pistil. 


168 THE FLOWER. 


gous with leaves; that the sepals are comparatively little, the 
petals more, and the reproductive organs much modified from the 
type, that is from the leaf of vegetation. This is simply what 
is meant by the proposition that all these organs are transformed 
or metamorphosed leaves. What would have been leaves, if 
the development had gone on as a vegetative branch, have in 
the blossom developed in other forms, adapted to other fune- 
tions. Linnzeus expressed this idea, along with other more 
speculative conceptions, dimly apprehended, by the phrase Vege- 
table 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 Goethe, 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 foliaceous and floral 
organs.* 


1 The contribution of Linnzus is on p. 301 of the Philosophia Botanica, 
1751; and all that is pertinent is in the following propositions : — 

Plumulam seminis sepius terminat aut flos aut gemma. 

Principium florum et foliorum idem est. 

Principium gemmarum et foliorum idem est. 

Gemma constat foliorum rudimentis. 

Perianthium sit ex connatis foliorum rudimentis. 

His dissertation, Prolepsis Plantarum, in Ameen. Acad. vi. (1760), added 
nothing but obscure speculations to the former comparatively clear 
statements. 

Kaspar Friedrich Wolff’s contribution is in his Theoria Generationis, 
mainly concerning animals, published in 1759, 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-scales or floral 
organs, as the case may be, through degenerescence or diminution of vege- 
tative force, which is renewed in the bud or in the seed. 

Johann Wolfgang Géthe’s Versuch die Metamorphose der Pflanzen zu 
erkliren was published in 1790, in 86 pages. For the translations and 
reproductions, see Pritzel, Thesaurus. To the French translation by Soret, 
with German text accompanying (Stuttgart, 1831), and also to that of Ch. 
Martens ((Euv. Hist. Nat. de Geethe, Paris, 1837), are joined the author’s inter- 
esting notes and anecdotes of later periods, down to 1831. The degenerescence 
by diminution of vegetative force with renewals by generation, propounded 
by Wolff, in Geethe’s essay takes the form of successive expansion and con- 
traction of organs. ; 

A. P. DeCandolle’s Théorie Elémentaire 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 wnity 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, ¢. e. they conform 
to phyllotaxy, 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. 

397. 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 by transitions or 


a second edition 1519; a third (revised by Alphonse DeCandolle), in 1844, 
is posthumous. ‘The Organographie Végétale, 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 
Geethe in 1790, and that both Wolff’s and Geethe’s were unknown to DeCan- 
dolle until after the publication of the second edition of the latter’s Theorie 
Elémentaire, 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 Reper, 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 
a oe and texture it is not rare to 

i V oes, meet with bracts which vie 
Wf oe with, or indeed surpass, pet- 
- als themselves in delicacy and 

> brightness ; ae in such cases 


— : x aes oe for attraction. 
Ee geazle Sage, Painted-Cup 
en oe # (Castilleia) , and the Poin- 
(Ae 
Ae Bee settia, with other Euphorbias 


us 


, OSS of the conservatories, are ex- 
i. = i \ 7 . 

iXisx/ amples of this. In the flowers 
~~ 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- 
phea, Fig. 318), 
there is a gradual 
transition from the 
sepals through the 
petals to stamens ; 
in Lilies and most 
lily-like flowers, se- 
pals are as brightly 
colored as_ petals, 
and commonly more 
or less combined 
318 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, be expected. 
308. Teratological Transitions and Changes. Teratology is 
the study of monstrosities. These in the vegetable kingdom 


FIG. 317. Cactus-flower (Mamillaria cespitosa), with bractlets, sepals, and petals 
passing into each other. 
FIG. 318. Series exhibiting transition from sepals to stamens in Nymphza odorata. 


ITS METAMORPHY. wie 


often elucidate the nature of organs.! The commonest of these 
changes belong to what was termed by Geethe 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.? 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- ,——~ 
z ie ~ 
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 Tératologie Végetale, 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 Engelmann, 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 formativus; 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 down the inner edge, and the margins of the pistil-leaf bearing leaves 
instead of ovules. 


172 THE FLOWER. 


buttercups.! In these the green hue of the centre of the rosette 
indicates a tendency to retrograde a step farther into sepals, or 
into a cluster of green leaves. This takes place in certain blos- 
soms of the Strawberry, the Rose, &c. Such production of 

‘‘oreen roses,” and the like, has been appropriately called 
chlorosis, ov by Masters chloranthy, from the change to green. 

309. 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 

ff) the latter may be compared with the leafy 

/ 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, 521), usually 

passing moreover, by prolification of the re- 

; ceptacle, into a leafy branch. 

ILO yA 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 
very common, and curious grades between the two are met with 
almost every spring. So also in the common Houseleek, 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 Jt must not be concluded that the supernumerary petals in all’such cases 
are reverted stamens, or stamens and pistils. Some are instances of abnormal 
pleiotaxy, 7.e. of the production of one or more additional ranks of petals 
(better deserving the name of double flower), with or without reversion of 
essential organs to flower-leaves. 

2 These trees are popularly supposed to bear fruit without blossoming; 
the reverted green petals being so inconspicuous that the flower is un- 
noticed. 


FIG. 320, 321. Green leaves from the centre of a blossom of Double-flowering Cherry, 
one still showing, by its partial involution and its style-like apex, that it is a reverted 
carpel, the other a small but well-formed leaf. 

FIG. 322. Leaf or leaflet of Bryophyllum, developing plantlets along the margins. 


ITS METAMORPHY. 133 


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 may be. 
It may resume yegeta- 
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. 524, 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 leafy 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 | eas 
flower-bud) makes its appearance in the eee he! 
axil of a petal or of a stamen; and it 
may be clearly inferred that the organ 


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 bearing 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 (Nymphza Lotus) is figured 
and described by Dr. Masters.! 

5314. In the application of morphological ideas to the elucida- 
tion of the flower, nothing should be assumed in regard to it 
which has not its proper counterpart and exemplar in the leaves 
and axis of vegetation. 


Section II. FrLorat Symmetry. 


315. The parts of a flower are symmetrically arranged around 
its axis.”, 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. 

316. Adopting the doctrine that the parts of the flower are 
homologous with leaves, the synimetry 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. 

Phyllody (called Phy/lomorphy by Morren, “rondescence by Engelmann) is 


the condition wherein true leaves are substituted for some other organs; 


i.e., where other organs are metamorphosed into green leaves. There is 
phyllody of pistils, ovules, filaments, anther, petals, sepals, &c. 

Sepalody, where other organs assume the appearance of green sepals. 

Petalody, where they assume the appearance of petals, as normally in 
Pinckneya and Calycophyllum, in which one calyx-lobe enlarges and becomes 
petal-like, and abnormally in Primroses where all the calyx-lobes imitate 
lobes of the corolla (this has been termed Calycanthemy) ; also of the stamens 
of common “ double flowers.” 

Staminody, where other organs develop into stamens. Cases of this as 
affecting pistils are referred to above: rarely sepals and petals are so affected. 

Pistillody, where other organs develop into pistils, which most rarely 
happens except with the stamens, as above mentioned. 

2 Tt is stated that Correa 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 Théorie Elementaire, and elaborated in detail by A. St. 
Hilaire in his Morphologie Végétale. 


FLORAL SYMMETRY. ae 


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 ¢somerous. 

317. A Symmetrical Flower is one in which the members of all 
the cycles (whorls or seeming whorls) are of the same number.? 
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 
tsomerous flower (meaning one with an equal number of mem- 
bers of all organs) is the same as symmetrical, 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 ; 7. 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 defined. 

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 “ simply 
symmetrical or monosymmetrical,” and those which can be symmetrically 
divided by two or more planes, “doubly symmetrical or polysymmetrical,” 
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, Nym- 
pheea for all but perLaps the sepals, many Cactacez 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 Hip- 
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, quaternary, quinary, senary, 
&¢., 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 

Monomerous, for the case of a flower of one member of each ; 

LDimerous, 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,' of six, or on the plan of six members to each 
circle. But, in Monocotyledons, so-called hexamerous blossoms 
are really trimerous, the sixes being double sets of three. 

523. Pattern Flowers. These should be symmetrical, regular, 

complete in all the parts and without ex- 
; \)/ cess or complication of these, and with- 


: * te Vig out any of the cohesions or adhesions 
u " {/ C = \ which may obscure the type, or render it 
3) &O less expressive of the idea that a flower 
c W ee consists of a series of circles or spirals 
= of modified leaves crowded on a short 

my = axis. Wherefore the illustration Fig. 307, 


with its diagram Fig. 308, may serve as a pattern pentamerous 
or quinary flower; and Fig. 326, with its diagram, Fig. 327, 


1 These may be shortly written 1-merous, 2-merous, 3-merous, and so on 
to 10-merous (decamerous), 12-merous (dodecamerous), &c. 


Fig. 326. Parts of a symmetrical trimerous flower (Tillea muscosa): @. calyx; 
6 corolla; ¢c. stamens; d. pistils. 327. Diagram of the same. 


FLORAL SYMMETRY. TUT 


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 
d-merous flower as that displayed 

in Fig. 328. 
ha 324. Diplostemonous 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.t As _ the 
petals alternate with 
the sepals, so the first 
series of stamens al- 
ternates with the pet- 
323 als, the second series 2 
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. Pl. Oudney, in-Denham and Clapperton Tray. 1826, reprinted in Ray 
Soe. ed. of Collected Works, i. 293). It is true that Brown declares the 
same of the pistils; but that isnot 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 andreecium 
or the blossom is said to be 

Tsostemonous 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 
pentamerous 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, ), 
a petal, c, a stamen, and d, a pistil, also shown, enlarged. 

FIG, 329. A pentamerous diplostemonous flower of Seduna. 


178 THE FLOWER. 


superposed to) the sepals, and the stamens of the inner series 
stand before the petals; as in the diagram, Fig. 331.7 
325. Flowers which completely 
exemplify their type or symmetry 
are rare, but most exhibit it more 
or less. Each natural order or 
group exhibits its own particular 
floral type, or modification of the 
common type.2 Some of these 
modifications do not at all affect 
the symmetry 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 plants 
331 of the same order or tribe, or of related 
orders. The symmetry 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 
writers term the stamens which are before the petals ep/peta/ous, those before 
the sepals episepalous ; but, as this prefix means upon, it is better to restrict 
these terms to cases of adnation of stamens to these respective parts of the 
_ perianth, and to distinguish as 

Antipetulous, those stamens which stand before petals, whether adnate or 
free, and 


Antisepalous, those which stand before 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. 

Parapeta/ous, for stamens which stand at each side of a petal, yet not 
necessarily before a sepal, as in many Rosacez. 

2 These particular types, with their modifications, are set forth in the 
characters or distinguishing marks of the orders, tribes, genera, &e. The 
best generally available illustrations of ordinal types are in Le Maout and 
Decaisne’s Traité Général de Botanique, and in Hooker’s English edition 
and revision, entitled A General System of Botany, Descriptive and Analyti- 
cal, London, 1873. The best morphological presentation is in Eichler’s 
Bliithendiagramme, &c. (Flower Diagrams, Constructed and Illustrated), 
Leipzig, 1875. 


FIG. 350. Opened flower of Trillium erectum. 331. Diagram of the same. 


a 


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. Varrous MoprricaTions 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 eyen in that one of the orders, the Crassulacez, 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; Tilleea (Fig. 326) is the 
same, but with the members symmetrically 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 the three remaining coalesce into one body up to the 


180 THE FLOWER. 


middle; Penthorum (Fig. 335, 336) has its five carpels coales- 
cent almost to the top, and usually loses its petals by abortion ; 
in Grammanthes and Cotyledon (Fig. 332-334), the sepals are 
coalescent into a cup and the petals into a deeper one, out of 


which the stamens appear to arise, these being adnate to the 
corolla. Symmetrical increase in’ the number of members of 
each circle is no proper deviation from type, at least in this 
family (in which flowers on the same plant sometimes vary from 
5-merous to 4-merous and 6-merous) ; and in Sempervivum (to 
which Houseleek belongs) these members are always more than 
five and sometimes as many as twenty in each circle. 


§ 2. REGULAR UNION OF SIMILAR PARTs. 


329. Coalescence, or the cohesion by the contiguous margins 
of parts of the same circle or constituent set of organs, is so fre- 
quent that few flowers are completely free from it. The last 
preceding figures show it in the gynecium and corolla. Fig. 
471-476 further illustrate it in the corolla, and in various degrees 
up to entire union; and Fig. 483-488 illustrate it in the andree- 
cium. ‘he technical terms gwhich coalescence calls for, and 
which are needful in botanical description, may be found under 
the account of the particular organ, and in the Glossary. Such 
growing together of contiguous members in the blossom is strictly 
paralleled by connate-perfoliate leaves of ordinary foliage (212, 
Fig. 215), where it more commonly occurs in upper leaves, and 
in bracts, which are still nearer the flower. 

330. It should now be hardly necessary to explain that the 
terms coalescence, cohesion, union, and the corresponding phrases 


FIG. 332. Flower of Grammanthes. 333. Flower of a Cotyledon. 334. The corolla 
laid open showing the two rows of stamens inserted on it. 335. The tive pistils of 
Penthorum, united. 336. A cross-section of the same. 


UNION OF PARTS. 181 


in the next paragraph, do not mean that the parts were once sepa- 
rate and have since united. That is true only of certain cases. The 
union is mostly congenital, equally so in the disks of foliage of 
the Honeysuckle (Fig. 215) and in the corolla of a Convolvulus. 
The lobes which answer to the tips of the constituent leaves of 
the cup or tube are usually first to appear in the forming bud, 
the undivided basal portion comes to view later. It might be 
more correct to say that the several leaves concerned have not 
isolated themselves as they grew. Accordingly, Dr. Masters 
would substitute for coalescence and adnate the term inseparate. 
But the common language of morphology needs no change, as it 
consistently proceeds on the idea, and the prevalent fact, that 
leaves are separate things, and that the tube, cup, or ‘* insepa- 
rate” base of a calyx or corolla, consists of a cértain number of 
these. It is no contradiction to this view that they developed 
in union.’ 


§ 3. UNION OF DISSIMILAR OR SUCCESSIVE Parts. 


331. Adnation is the most appropriate term to denote the organic 
and congenital cohesion or consolidation of different circles, the 


1 If it were *eriously proposed to change the language of descriptive 
botany in this regard, consistency would require its total reconstruction, with 
the abolition of all such terms as cleft, parted, &c.; for the structures in 
question are no more cleft than they are united. While these convenient 
and long-familiar terms are continued in use (as they surely will be), although 
quite contrary to literal fact, it cannot be amiss to continue those, such as 
connate, adnate, coalescent, &c., which imply and suggest the fundamental fact 
in the structure of phenogamous and the higher cryptogamous plants, viz. 
that leaves are normally unconnected organs. : 

Whether fusion or separation is the more complex condition, and therefore 
indicativé of higher rank, is a question of a different order. It is argued 
that the fusion or lack of separation is an arrest of development, and there- 
fore an indication of low rank or less perfection than the contrary. Buta 
phylogenetic view of the whole case may reverse this conclusion as respects 
the blossom. ‘The course of development from thallus and frond to distinct 
foliage on an axis, from little to full differentiation, is clearly a rise in 
rank, as also is the differentiation of fdliage into ordinary leaves, petals, 
stamens, and pistils. But there is as much differentiation in the flower 
of a Convolvulus as of a Ranunculus, and more in that of a Salvia, a 
Lobelia, and an Orchis. In all such flowers, the combination, the irregu- 
larity, and the diversification in many cases of the members of the same circle, 
all indicate complexity, greater specialization, and therefore higher rank. 
The production of leaves distinct from the axis is one step in the ascending 
scale: such specializations and combinations of these as occur in flowers are 
higher steps ; and the most specialized, complex, and therefore highest in 
rank are complete, corolliferous, irregular flowers, with a definite number of 
members, and these combined in view of the adaptations by which the ends 
of fertilization and fructification are best subserved. 


182 THE FLOWER. 


apparent growing of one part on or out of another, —as of 
the corolla out of the calyx, the stamens out of the corolla, or 
all of them out of the pistil. This disguises the real origin 
of the floral organs from the receptacle or axis, in successive 
series, one within or above the other. Organs in this condition 
are also and rightly said to be Connate (born united) ; but, as 
this term is equally applicable to the coalescence of members of 
the same circle, the word Adnate is preferable, as applying to 
the present case only. Adnation is heterogeneous organic co- 
hesion or adhesion: coa- 
lescence is homogeneous 
cohesion or union. 

332. Adnation occurs in 
very various degrees, and 
affects either some or all the 
organs of the flower. Its 
consideration introduces into 
terminology several peculiar 
terms, which may here be 
defined in advance. Three 
of them, introduced and 
prominently employed by 
Jussieu, depend upon the 
degree of adnation, or the 
absence of it, viz. :— 

Hypogynous (literally be- 
neath pistil) , applied to parts 
which are inserted (7. e. are 
borne) on the receptacle of 
the flower, as in Fig. 336. 
This is the absence of 
adnation, or the condition 
which corresponds with the 
unmodified type. 

Perigynous (around the 
pistil) implies an adnation 
which carries up the inser- 
tion of parts (which always means apparent origin or place of 
attachment) to some distance above or away from the recep- 


20Q 
538 


FIG. 336%. Vertical section of a flower of the Common Flax, showing the normal or 
hypogynous insertion of parts upon the torus or receptacle. 

FIG. 237. Vertical section of a flower of the Cherry, to show the perigynous insertion, 
or adnation to the calyx, of the petals and stamens. 

FIG. 338. Similar section of the flower of the Purslane, showing an adnation of all 
the parts with the lower half of the ovary. 


UNION OF DISSIMILAR PARTS. 183 


tacle, so commonly placing this insertion around instead of 
beneath the pistil; whence the name. The perigyny may be, 
as the figures show, merely 
the adnation of petals and 
stamens to calyx, the calyx 
remaining hypogynous, as in 
Fig. 337; or else the adna- 
tion of the calyx, involving 
the other organs, to the lower 
part of the ovary, as in Fig. 
3358, or up to the summit of 
the ovary, while the petals 
and stamens are adnate still further to the calyx, as in Fig. 339. 
The latter passes into what is called 

Epigynous (on the pistil), where the adnation is complete to 
the very top of the ovary, and none beyond it, as in Fig. 340, 
541. Yet here the parts so termed are not really on the ovary, 
except where an epigynous disk (394) actually surmcunts it. 


333. Adnation brings some other terms into use in botanical 
descriptions, especially those of superior and inferior. In this 
connection, these words (in Latin taking the form of superus and 
inferus) denote the position in respect to each other of ovary and 
floral envelopes, — not the morphological, but the apparent posi- 
tion or place of origin. Thus, in Fig. 836 and in 337, the calyx 
is ¢nferior, or in other words the ovary superior. Here real and 
apparent origin agree, this being the normal condition, which 
is otherwise expressed by saying that the parts are free, 7. e. free 
from all adnation of one to the other. But, in Fig. 339-341, the 


FIG. 339. Similar section of a flower of Hawthorn, showing complete adnation to 
the summit of the ovary and of the other parts beyond. 

FIG. 340. Vertical section of a Cranberry-flower, and 341, of flower of Aralia 
nudicaulis, with so-called epigynous insertion of calyx, corolla, and stamens; the calyx 
of the latter completely consolidated with the surface of the ovary, or its limb 
obsolete. 


154 THE FLOWER. 


ovary is said to be inferior and the calyx superior, the calyx 
and other parts, in consequence of the adnation of its lower part, 
seeming to rise from the summit of the ovary. 

334. Adnation of floral envelopes to pistil rarely extends 
beyond the ovary; yet, in species of Iris having a tube to 
the perianth, this tube is commonly adnate for most of its length 
to the style. But when the calyx has its tube or portion with 
united sepals prolonged, the petals and the stamens are usually 
adnate more or less to it, 7.e. are inserted on the calyx. And, 
when the petals are united and prolonged into a tube, the sta- 
mens, being within the corolla, are commonly adnate to or 
inserted upon this. . 

335. No one doubts that the view is a true one which repre- 
sents the perianth-tube as adnate to the style in Iris, petals and 
stamens as adnate to calyx in the Cherry (Fig. 337), stamens 
as adnate to base of corolla in Fig. 334, and a long way farther 
in Phlox, &c. That the calyx is similarly adnate to the ovary 
is nearly demonstrable in certain cases. 

336. But, as the lower portion of a pear is undoubtedly recep- 
tacle, or rather the enlarged extremity of the flower-stalk, as in a 
rose at least a portion of the hip is receptacle, as the tube of the 
flower in a Cereus or other Cactacea has all the external char- 
acters and development of a branch, so it is most probable that 
in many cases the supposed calyx-tube adnate to an inferior 
ovary is partly or wholly a hollowed receptacle (in the manner 
of a Fig-fruit) ; that is, a cup-shaped or goblet-shaped develop- 
ment of the base of the floral axis. This would bring the case 
under § 7. (326, 495.) 


§ 4. IRREGULARITY OF SIMILAR PARTS. 


337. Irregularity, or inequality in form or in union of mem- 
bers of a circle, is extremely common, either with or without 
numerical symmetry. One or two examples may suffice. 

338. Irregular flowers with symmetrical perfection, except in 
the gyneecium, are well seen in the Pea Family, to which belongs 
the kind of corolla called Papilionaceous, from some imagined 
resemblance to a butterfly. (Fig. 342-344.) This flower is 
5-merous throughout, has the full complement of stamens (10, or 
two sets), but the gynecium reduced to a single simple pistil. 
The striking irregularity is in the corolla, the petals of which 
bear distinguishing names: the posterior and larger one, exter- 
nal in the bud, is the Vexititum or Stanparp (Fig. 344, a); 


INEQUALITY OF SIMILAR PARTS. 185 


the two lateral next and under the standard, ALa& or Wines (4) ; 
the two anterior, covered by the wings and partly cohering to 


O-4 


form a prow-shaped body (c), the Carma or Kret. The calyx 
is slightly irregular by unequal union, the two upper sepals 
united higher than the 
other three. The sta- 
mens are much more 
coalescent,but with an 
irregularity, nine com- 
bined by the lower part 
of their filaments, and 
one (the posterior) 
separate. (Fig. 345.) a 

339. The plan and * stad 
floral symmetry in the Locust-blossom and 4; 
its relatives are little obscured by the irregu- 
larities and the coalescence, hardly more so 
than in the plainer flower of its relative, 
Baptisia (Fig. 347, 348), in which the petals are somewhat 
alike, and the ten stamens are distinct or unconnected. Only 
the calyx is more irregular, by the union of the two posterior 
sepals almost to the tip. (Fig. 348.) 


548 


FIG. 342. Diagram of flower of the Locust, Robinia Pseudacacia: a. axis of inflores- 
cence; 6. bract; first circle of 5, calyx; five remaining pieces, corolla; next anthers, 
10 in number; in the centre a single simple pistil. 343 Front view of Locust-flower 
showing only the corolla. 344. This corolla displayed. 

FIG. 345. Andreecium of the Locust, nine stamens coalescent, one distinct. 346 
Same of a Lupine, all ten filaments coalescent below into a closed tube. 

FIG. 3!7. Calyx and corolla of Baptisia australis. 348. Same with petals fallen, 
showing ten distinct stamens and tip of the style. 


186 THE FLOWER. 


340. But in a Lupine-blossom, of equally near relationship, 
a casual observer might fail to recognize the very same type, 
although disguised only by cohesions. For while the two pos- 
terior sepals are united to the tip on one side of the blossom, the 
three others are similarly united into 
one body on the anterior side, giving 
the appearance of two sepals instead 
of five: in the corolla, the two keel- 
petals are more strictly united into a 
slender scythe-shaped or sickle-shaped 
body; so that the petals might with 
the unwary pass for four: in the 
andreecium, the coalescence includes all 
ten stamens (Fig. 346), which is an 
approach to regularity. 

341. The 5-merous symmetry of the 
Violet-blossom is complete until the 
gyneecium is reached (but with only 
one circle of stamens) ; the main irregu- 
larity of the perianth is in the anterior 
petal, with its nectariferious sac at base 
(Fig. 349-351) ; the two stamens near- 
est this send into the sac curious appendages, which the other 
three do not possess; the gynecium is composed of three car- 

3) pels coalescent into one compound ovary in a 
manner hereafter explained. In Antirrhinum 
yy and Linaria (Fig. 480, 481), there is a similar 


UN 
(j irregularity accompanying coalescence of the 
petals, the anterior one being extended at base 


Me WW into a nectariferous sac or hollow spur.! The 
/ flower of a Lobelia (Fig. 488) has the same 


See numerical plan and symmetry as that of Viola 

351 (except that the gyncecium is dimerous) ; but 

the members are adnate below and coalescent above, and the 

corolla is irregular through unequal coalescence of the five petals, 
and the absence of coalescence down one side. 


1 PeLoria is a name given by Linnzus to an occasional monostrosity of 
these flowers (imitated in sundry others), in which the base of every petal, or 
answering part of the corolla, is prolonged downward into a sac or spur. 
The sac is, morphologically considered, a departure from normal regularity : 
in the monster, symmetrical regularity is restored by the development of four 
more sacs. 

FIG. 349. Flower of Viola sagittata. 350. Its sepals and petals displayed. 351. 


Diagram of a Violet-blossom, from Eichler, with bract or subtenling leaf (below), a pair 
of bractlets (lateral), and 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 Cirele. 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 organs. 
In Amorpha, of the same family, these four 
petals are gone, leaving no trace, reducing 
the corolla to a single petal. (Fig. 352, 355.) 
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 andreecium is straight, all irregularity thus 
disappearing through suppression. om 

344. Delphinium or Larkspur and Aconite or Monkshood 
furnish good examples of flowers in which irregularity is aecom- 
panied by more or less abortion. The calyx of the Larkspur 
(Fig. 354-356) is irregular by 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, 


188 THE FLOWER. 


four in number, and these 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. 355. The five sepals (outer circle) and the four 
petals (inner circle) displayed. 356. Ground-plan of the calyx and corolla. 

FIG. 357. Flower of an Aconite or Monkshood. 358. The five 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 dotted line) is vacant, this petal being suppressed, 
thereby rendering the blossom unsymmetrical. In Aconite 
(Fig. 857-359), 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 Linneeus 
NEcTARIES, a somewhat misleading name, as_ they 
are no more devoted to the secretion of nectar than 
ordinary petals or other partsare. In these flowers, 
moreover, the stamens are much increased in number. 

345. Analogous abortion of some of the stamens, along with 
a particular irregularity of the perianth, especially of the corolla, 
characterizes a series of natural orders with coalescent petals.! 
These flowers are all on the 5-merous plan (except that the 
gyneecium 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 
Scrophulariacee (Snapdragon, Pentstemon, Mimulus, &c.), Orobanchacee 
(Beech-drops), Lentibulacee (Bladderwort), Gesneracee (Gloxinia), Big- 
noniacez (Trumpet Creeper, Catalpa), Pedaliacew (Martynia), Acanthacee, 
Labiate (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. 563) : 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 gyneecium 
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 pentacarpellary genus 
is a monstrosity. 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 the type, and indeed are sometimes present in 
that blossom’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 

Incomplete, in which any one or more of the four kinds of organs 
is wanting, whatever these may be; 

Apetalous, when the corolla or inner perianth is wanting ; 

Monochlamydeous, where the perianth is simple instead of 


FIG. 361. Corolla of Gerardia purpurea laid open, with the four stamens, the place 
which the fifth should occupy indicated by a cross. 

FIG. 362. Corolla of Pentstemon grandiflorus laid open, with its four stamens, and 
a sterile filament in the place of the fifth stamen. 

FIG, 363. Corolla of Catalpa laid open, with two perfect stamens and tlie vestiges 
of three abortive ones. 


DISAPPEARANCE OF PARTS. 191 


double, in which case the wanting set is generally (but not quite 
always) 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 ; 

Monecious (of one household), when stamens and pistils oc- 
cupy different flowers on the same plant ; 

Diecious (of two households), when they occupy different 
flowers on different plants ; 

Polygamous, when the saine species bears both unisexual and 
bisexual or hermaphrodite flowers. This may occur in various 
ways, from the greater or less abortion of either sex, either on 
the same or on separate individual plants ;— as Moneciously or 
Dieciously Polygamous, according to the tendency to become either 
monececious or dicecious. Recently Darwin has well distinguished 
the case of 

Gyno-diecious, where the flowers on separate individuals are 
some hermaphrodite and some female, but none male only ; and 
Andro-diecious, 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. (556, 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 astaminate one; a fertile flower, one which 
is pistillate, if not also hermaphrodite. 

348. Suppressed Perianth. Almost universally, when the peri- 
anth is reduced to asingle 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 difference 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. 

5349. Yet the only perianth obvi- 
ously present may be corolla, as when 


to the ovary and its border or lobes obsolete or wanting.’ Aralia 
nudicaulis (Fig. 541) is an instance, likewise many Umbellifere, 
some species of Fedia or Valerianella, the fertile flowers of Nyssa, 
and those Compositze which have no pappus. For Pappus, 
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. (644, Fig. 631-633.) 
When the obliteration is complete, as in Mayweed (Fig. 630), 
in some species of Coreopsis, &c., the corolla seems to be simply 
continuous with the apex of the ovary. A comparison with 
related forms reveals the real state of things.? 

350. So also in Hippuris, 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 
calyx, and near inspection detects its border. 

351. Both calyx and corolla are really want- 
ing in the otherwise complete and perfect (symmetrical and 


1 In the flowers of the two common species of Prickly Ash (Zanthoxy- 
lum) of the Atlantic United States, one has a double, the other a single 
perianth (as shown in Gray, Gen. Illustr. ii. 148, t. 156): the position of the 
stamens gives a presumption that the missing circle of the latter is the calyx ; 
yet it may be otherwise explained. In Santalacee there are some grounds 
for suspecting that the simple perianth, although opposite the stamens, is 
corolla; and the foliaceous sepal-lobes of the female flowers of Buckleya 
woud confirm this, if these are true sepals rather than adnate bracts. 

2 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. 


FIG. 364. Flower of Anemone Pennsylvanica; apetalous, the calyx petaloi I. 
FIG. 365. Achlamydeous flower of Lizard’s-tail (Saururus cernuus), magnided. 


the calyx has its tube wholly adnate ~ 


2 


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. 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 Andrecium or Gynecium. This occurs with- 
out or along with suppression in the perianth. In cases of the 
former, vestiges of 370 _. oon 
the aborted organs WW, 
often remain to sig- , 
nify the exact nature 
of the loss. Sepa- 
ration of the sexes 
(monecious, diceci- 
ous, &c.) is the re- 
sult of such suppres- 
sion. In Menisper- 
mum (Fig.368, 369), 
this is accompanied 
by an actual doub- 
ling of both calyx 
and corolla. ‘The 
dicecious flowers of 
Smilax 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. Andreecium and gynecium; the former 
of 5 perfect stamens, alternating with 5 rudiments of a second set. 

FIG. 368, 369. Dicecious 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. 


{94 THE FLOWER. 


354. 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 
in Willows, the flowers of which are all achlamydeous 
and diecivus. (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. 
370-372) 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. 

359. Still more 
simplified flowers, 
but more difficult 
to comprehend, are 
those of Euphorbia, 
or Spurge. These 
are in fact moneeci- 
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 flower. (Fig. 376, 377.) This umbel-like 


FIG. 374. A separate staminate flower of Salix purpurea, with the stamens coa- 
lescent (monadelphous and syngenesious), so as to appear like a single one. 

FIG. 375. Flowering branch of Euphorbia corollata. 376. Calyx-like involucre 
divided lengthwise, showing the staminate flowers around a pistillate flower (a). 377. A 
more magnified staminate flower detached with its bract, a; its peduncle or pedicel 5, 
surmounted by the solitary stamen,c. 378. Pistil in fruit, cut across, showing the 
three one-seeded carpels of which it is composed. 


SUPERPOSITION OF SUCCESSIVE PARTS. 199 


flower-cluster 1s 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 they exactly imitate petals. Here, then, is 
a whole cluster of extremely simplified flowers, taking on the 
guise of and practically behaving like a single flower, the invo- 
lucre serving as calyx and corolla ; the one-stamened male flowers 
collectively imitating the andreecium 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 world, 
gives proof that.this interpretation is the true one. 

356. Suppression of beth Andrecium and Gynecium. ‘This 
occurs in what are termed Neutral Flowers (347), such as are 
conspicuous at the margin of the cymes of Hydrangea (Fig. 295) 
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 fifth 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 Fig. 369: in the male blossom this is obscured in the andree- 
cium (Fig. 368) by a multiplication of the stamens.’ The 
same thing occurs in the perianth and bracts of certain Clusiacee, 
in which the members counted as in fours are superposed, and 
in some of which the double dimerous arrangement with apparent 
anteposition extends through the corolla; while, in other closely 
related flowers, the corolla changes to simply tetramerous and to 
alternation with the preceding four sepals. This passes, in the 
same family and in the allied Ternstroeemiacez, into 

360. Superposition by Spirals, as where five 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.” The flower 
of Camellia is continuously on the spiral plan up to the gyne- 
cium ; but upon one which, from the bracts onward, rises from the 
4 to the 2 and 2 order or higher, throwing the petals of the rosette 
in a full-double fiower into numerous more or less conspicuous 
vertical ranks. 

361. Anteposition in the Andrecium. It is in the andreecium 
that 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. (524. ) 

362. With Isostemony. Vitis (Fig. 879-381), also Rhamnus 
(Fig. 415, 416), and the whole Grape and Buckthorn families of 


1 In Columbine (Aquilegia), multiplication of the stamens in successively 
alternating 5-merous whorls similarly brings the andreecium into ten ranks; 
so, when these stamens in double flowers are transformed into hollow-spurred 
petals, these are set one into another in ten vertical ranks. 

2 Gen. Illustr. ii. t.140. But the petals alternate with the sepals in the 
ordinary manner of the flower, though their strong quincuncial imbrication 
suggests the spiral arrangement. 


e = 
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 = 8% 580 
the receptacle, alternate with and inside 
the stamens: there is no good reason to 
suppose that they answer to a second 
row of stamens. Allisostemonous Por- 
tulacaceze 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 Plumbaginacez, Primu- 
laceze, the related Myrsinaceze, and in 
most Sapotaceze, in the latter usually 
with some complications. a 

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.’ Observa- 
tion supplies no vestige of proof of it in Rhamnacez and Vitacez ; 
but, in the group of related orders to which the Primulacez 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 Sapotacez, while Chrysophyllum 
has in these respects just the structure of Primulacez, and 
Sideroxylon that of Samolus, Isonandra 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 Fichler, 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 hypothesis, there is no 
andreecial circle in such blossoms, or only vestiges of one, but the petals have 
supplied the deficiency by a supernumerary production of theirown! The 
more plausible hypothesis of Braun, 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. 


. 
198 THE FLOWER. 


completing the symmetry of the blossom and 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 androecium 
of a typical flower should consist of two circles of stamens. 
(324.) ‘The only serious objections to this explanation rise out of 
the difficulty 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, is also itself very 
commonly attended by anteposition. In normal or Direct 
Diplostemony, — 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 Obdiplostemony. This condition prevails, more or 
less evidently, in Ericacez, Geraniaceze, Zygophyllacez, Rutacez, 
Saxifragacex, Crassulacee, Onagracez, &c. (but in some of these 
with exceptions of direct diplostemony) ; also, accompanied 
by a peculiar multiplication of members (380), in Malvacez, 
Sterculiaceze, and Tiliacee. The explanation is difficult. 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. Diagram of pattern flower with direct diplostemony. 383. Diagram 
of similar flower with obdiplostemony. Both from Eichler’s Bliithendiagramme. 


* 


SUPERPOSITION OF SUCCESSIVE PARTS. 199 


(whether adnate 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 ae 
antisepalous stamens the third floral circle 7%, 2~ 7" <\ 
or the only truly andrecial one, and sym- / “Be \ 
metrically alternate with the petals on the one ( Se 2) 
hand and the carpels on the other. The NY G 2 o} 
second hypothesis conceives that there is a \\So ee oy 
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. 354), were plausibly 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 Celakowsky, 
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.? 


384 


1 Jn 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 Caryophyilacee) 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, @. e. sepals, 
petals, greenish bodies called glands, antipetalous stamens, antisepalous stamens, 
carpels. 


200 THE FLOWER. 


366. The case of stamens in a cluster before the petals is a 
complication of either 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 modilications of the type. It occurs in two 
ways: 1st, by an increased number of circles or turns of spirals 
in the flower, which is Regular Multiplication ; 2d, by the pro- 
duction of two or three or of many organs in the normal place 
of one, Chorisis or Deduplication. ‘The first does not alter the 
normal symmetry of the blossom, although it may render it dif- 
ficult or impossible to trace or demonstrate it. The second 
apparently disturbs, or at least disguises, floral symmetry. 
Either may be definite, or of a constant and comparatively 
small number; or c¢ndefinite, when too numerous for ready 
counting, or inconstant, as the higher numbers are apt to be. 

368. Regular Multiplication, or Augmentation of flural circles 
or spirals, may affect any or all the four organs, but most com- 
monly the andreecium. When the perianth is much increased 
in the number of its members, the distinction between calyx and 
corolla, or even between bracts and corolla, is apt to disappear, 
as in most Cactaceous flowers (Fig. 317), Nelumbium, Calycan- 
thus, &c. In these and similar cases, the members of the perianth 
are prone to take a spiral instead of cyclic arrangement ; and this 


and in obdiplostemony. Along with the lack of clear analogy to support 
St. Hilaire’s hypothesis of transverse deduplication, the similar orientation 
of the vascular bundles in the petal and the stamen before it must, as 
Celakowsky insists, be good evidence that these represent independent 
leaves, and not superposed portions of one. 

The main objection to the second hypothesis (that of a suppressed 
circle outside of the antipetalous stamens) is that this missing circle, 
whether of petals or stamens, is not actually met with in any nearly re- 
lated forms (for in Monsonia the fifteen stamens are otherwise explained) ; 
also that there are transitions, as above mentioned, between obdiplostemony 
and direct diplostemony. To Braun’s theory that the glands behind the 
antisepalous stamens in true Geraniacee answer to suppressed phylla, 
Eichler objects that these are present behind all ten stamens in Oxalidee ; 
also that all are wanting when the oflice of nectar-secretion, which they sub- 
serve, is undertaken by some other part of the flower, as by the calyx-spur 
in Pelargonium and Tropeolum. 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 Anonacez, Ranunculus, 
Anemone, and the like. But in Aguilegia, where the number five 
is fixed in the perianth, the cyclic arrangement with alternation 
of whorls prevails throughout 

369. The definite angmentation of calyx and corolla by the 
production of one additional whorl of each, and the seeming 
anteposition which comes of it when the andrecium remains 
simply diplostemonous (in the manner of the Berberidacee, 
Menispermaceze, &¢c., 359) has already been explained. 

370. Similar increase to two whorls affecting the corolla only 
characterizes Anonacezr, Magnoliaceze, Papaveraceze, and Fuma- 
riacee. In all but the last order, this is accompanied by indefi- 
nitely multiplied stamens, and mostly by an increased number of 
earpels. In Fumariacez, which has dimerous flowers, there is a 
diminution by the suppression in most cases of half the normal 
andreecium, 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 Rosacez, but is not peculiar to it.? 
The simplest case, but a rare one, is seen in the 10-stamened vari- 
ety of some Hawthorns, as occasionally in Cratzegus 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-stamened 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 Pomez and many Potentillez, 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.” 


1 It was first clearly described by Dr. A. Dickson, in Trans. Bot. Soc. 
Edinb. viii. 468, and Seemann’s Jour. Bot. iv. 473 (1866). 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. Chorisis or Deduplication. Both these terms, and the 
ideas which they denote, originated with Dunal, but were first 
expounded by Moquin-Tandon.’ The first word is Greek for a 
separating or separation. The second is a translation of Dunal’s 
French word dédoublement (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. Hilaire) 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 flower, 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 are 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. It 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. Chorisis is complete when the parts concerned are dis- 
tinct or separate to the very insertion, as in the stamen-clusters of 
Hypericum. 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 
deduplications 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 antisepalous stamen, divided by chorisis (sometimes 
incompletely) and much separated, is not improbable. The other tenable 
explanation (which may be harmonized 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 asa whole. This makes them para- 
petalous, and at the same time brings them under Hofmeister’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 Sapotacez (in Trimen’s Jour. Bot. 1878). 
The inner circles are there sometimes 5-merous after the primitive type, 
sometimes 10-merous in regular alternation to the preceding circles. 

1 Moquin-Tandon, Essai des Dédoublemens, &c., Montpellier, 1826; Con- 
sidérations sur les Irrégularités de la Corolle, &c., in Ann. Sci. Nat. xxvii. 
237, 1832; Tératologie Végétale, 337. Dumnal, Essai sur les Vacciniées, 
1819, cited by Moquin (some pages printed, but never published) ; Considé- 
rations sur la Nature et les Rapports de quelques-uns des Organes de la 
Fleur, 1829. The next botanist to develop it was St. Hilaire, Morphologie 
Végétale, 1841. 


CHORISIS OR DEDUPLICATION. 203 


such sessile palmately compound leaves as those of some species 
of Aspalathus. It is incomplete when division does not extend 
to the base; as in Fig. 387, 395. 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 symmetry, whatever it be. 

374. St. Hilaire distinguished two kinds of deduplication; viz., 
collateral when the members stand side by side, and parallel 
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 may 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 Hypericum 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 at pe 
three, as is shown in the diagram, Fig. 386. 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 phulanr 
(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 way 


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. 


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 
of the inner series, trebled by chorisis, and this chorisis incom- 
plete, because it has not quite divided the filament into three. 
In Hypericum, the glands are completely suppressed, each pha- 
lanx is almost or quite divided into a cluster, either of about 
three stamens each, as in H. Sarothra, or of a few more (in H. 
mutilum and H. Canadense), or of an indefinite number, as in 
the common St. Johnsworts. Then in some other species (as 
in our H. pyramidatum) the carpels and the stamen-clusters rise 
to five, realizing complete pentamerous 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. Fumariacex, the 
Fumitory family, may 
furnish the next illus- 
tration. The flower is 
on the plan of two 
(dimerous) throughout. 


ZS 


S- 
oe 

392 393 
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 Cucullaria (Dutchman’s Breeches), ascape in flower and a leaf, 
severed from the singular bulb (formed of the enlarged bases of petioles). 389. Detached 
flower, of natural size, showing also the pair of bractlets on the pedicel. 390. Same 
with parts displayed, and 391, inner petals placed above. 392. Diagram of flower of 
Dicentra or Adlumia, from a section across the summit. 393. One of the phalanges of 
stamens of Adlumia; upper part only. 


CHORISIS OR DEDUPLICATION. 2O5 


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 asin 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 Adlumia (Fig. 393) and Corydalis 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 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 Fumariacez, or two in Crucifere, 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 development 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 THE FLOWER. 


378. The obvious relationship of Cruciferee to Fumariacee, 
their agreement in the rare peculiarity of having the two carpels 
595 side by side instead of fore and aft 
(median), and the characteristic 
anomaly which the andreecium pre- 
sents (7. 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, &c.), the pistil is 
of two carpels, right and left ; alter- 
nate with these is a pair of stamens 
on the side next the axis, matched 
by another pair on the opposite 
side of the pistil, the four longer 
and interior stamens; alternate 
with these, and lower in insertion, 
397 a single stamen on each side ; next, 
four petals, of somewhat various overlapping in eestivation, 
which essentially alternate with the two single stamens and the 
two pairs; lastly, four sepals, alternating with the four petals as 
a whole, the anterior and posterior overlapping the lateral ones 
in the bud. Now the median (7. e. the anterior and posterior) 
pairs of stamens occasionally have their contiguous filaments 
conjoined, as in Fig. 397. If this were at all constant, the 
inference would undoubtedly be that the case is one of chorisis, 
and that the flower as to its essential organs is dimerous. ‘This 
is apparently the best explanation to be given. It assumes that 
the chorisis is normally complete in the andreecium of Cruciferee, 
instead of incomplete, as in Fumariaceze.t And this view is 
confirmed by the fact that the median stamens are simple and 


1 The hypothesis here adopted, as to the andrecium, is that of Steinheil 
(1839), and of Eichler (in Flora, 1865, 1872, and Bliithend. ii. 200), replacing 
that of Kunth, 1833, &c., employed in former editions. The rejected view 
makes the flower 4-merous up to the pistil, and the stamens all of one circle. 
alternating with the four petals, the median stamens (as in our view) doubled 
by chorisis. Krause and Wretschko (cited as above by Eichler) would 
have the floral circles 2-merous and 4-merous by turns; the calyx of two 
2-merous circles (which it plainly is); the corolla of one 4-merous circle 


FIG. 394. A cruciferous flower. 395. Diagram of such a flower, with position of 
axis marked aboveit. 396. Tetradynamous stamens and the pistil. 397. A common 
monstrosity of the same, two of the four inner stamens combined into a common 
2-antheriferous body. 


CHORISIS OR DEDUPLICATION. 207 


single in Senebiera 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 Fumariaceze. 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 Crucifere, 
the blossom in the two orders would accord in having a 2-merous 
three-whorled perianth, the first and third whorls median ;? as 
also in the dimerous androecium, the first whorl of which is 
lateral. ‘The difference is that in Fumariacez the two members 
of the first whorl of stamens augment by chorisis into three, and 
the second is wanting, or is present only in Hypecoum ; while in 
Cruciferze the first whorl is simple (of the two short stamens), and 
the second is doubled. In Fumariacez only the first whorl of the 
perianth counts as calyx, and 
the corolla is of two whorls ; 
in Cruciferze, the first and 
second whorls are calyx, the 
inner sepals answering to the 
outer petals of Fumariacee. 

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 


399 


alternating with the calyx-members as a whole; the short stamens following 
as a 2-merous circle; then the long stamens as a 4merous circle; lastly the 
2-merous gynecium. 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, in 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. 


298 THE FLOWER. 


in each cluster with which the stamens cohere. The explanation 
by chorisis is that each cluster, petal-like body included, is a 
multiplication of one stamen. The diagram (Fig. 398) 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 
to the petals will take the same explanation as that of Rhamnus, 
Vitis, &c. (Fig. 363.) That the andreecium is here composed 
of the inner circle merely is partly confirmed by the alternation 
of the carpels with the clusters. According to Duchartre,’ the 
development of the andreecium 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 Sidaleea, as is most strikingly shown in the Californian 
S. diploscypha, has two series of stamens, the outer (answering 
to those of Malva and its relatives) in five membranaceous 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.? 

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. 3, iv. 123. Duchartre and 


others who draw freely upon median chorisis to explain anteposition, and 
consider that congenital union proves it, take the phalanges in these cases, 
like the single stamens in Vitis, to be an inner part of the petal itself. But 
this view appears to have had its day. 

2 Gray, Gen. Illustr. ii. 44, 57, 75-82. The position of the carpels before 
the petals in Pavonia 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. 


FIG. 400. A petal of Parnassia Caroliniana, with a triple staminodium before it. 
FIG. 4091, Diagram of the flower of Parnassia Caroliniana. 


OUTGROWTHS. 209 


alternate with the petals forms the effective andreecium. 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.* Wherefore, if 
these stamen-like bodies really represent a circle 
of the andrecium, it must be the inner one; and 
that is the more probable view. 

382. Multiplication by chorisis in the gyncecium 
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 
Malvacez, 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 Zrichomes (Trichoma, pl. trichomata), that is 
structures of which hairs are the type. They 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 conveniently illustrated under Vegetable Anatomy as 


1 Wichler in FI. Brasil., Sauvagesiace, & Bliithend. ii. 424; Drude in Lin- 
nea, xxxix. 239. Eichler refers to this as a confirmation of Celakowsky’s 
explanation of obdiplostemony by posterior displacement. (365.) 


FIG. 402. Pistil of Drosera filiformis with tricarpellary ovary (transversely divided), 
and six styles, i. ¢. three, and each two-parted. 


210 THE FLOWER. 


respects 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 may 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 trichomes. 
Examples of this are familiar in the pappus (answering to limb 
of the calyx) of many Compositz, and in the bristles which 
answer to perianth in many Cyperacez. The scarious stipules 
of Paronychia and of Potamogeton, the ligule of Grasses, and 
even the corolla in Plantago, are equally reduced to mere cellular 
tissue. So that the structural difference between trichomes and 
ouigrowths* is not at all absolute, and the morphological distine- 
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 Caryophyllacez, 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 Sapindaceze, 
these ligular 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 Emergenzen of the Germans, the 
Epiblastema of Warming, &c. For the development and discussion of this 
subject, see Warming, in Kjébenhavn Vidensk. Meddel. 1872, and a larger 
treatise on Ramification in Phanerogams, Copenhagen, 1872. Also, Uhl- 
worm in Bot. Zeit. 1873; Celakowsky in Flora, 1874; and Eichler’s note on 
Emergenzen in Bliithendiagramme, i. 48. 


FIG. 403. Petal of Silene Pennsylvanica, with its crown. 
FIG. 404. Flower of Passiflora ce@rulea, reduced in size. 


FORMS OF THE RECEPTACLE. 211 


are often more complicated in structure. They are always on 
the inner face, and are commonly two-lobed or parted. 

386. Similar stamineal appendages are well known in Cuscuta 
(Dodder), in Larrea (Fig. 405) and other Zygophyl- 
laceze, and less conspicuously in Gaura. 

387. To extend to them the name of LictLte may 
not be amiss, whether they are regarded as mere 
outgrowths of floral leaves, without further morpho- 
logical relations, or whether they be, at least some- 
times, interpreted as the homologue of intrapetiolar 
stipules, as their ordinarily two-cleft form, and their 
coincidence in Erythroxylum with an intrapetiolar two-cleft stipule 
suggest. 


§ 9. Forms or THE Torus OR RECEPTACLE. 


388. Torus is the more specific and proper name, RECEPTACLE 
is the more usual. (303.) A normal receptacle of the flower 
would be that of Fig. 316, the apex of the flower-stalk somewhat 
enlarged, roundish or depressed, and with surface mainly cov- 
ered by the insertion of the several organs; the Be inter- 
nodes which it potentially contains being 
undeveloped. As the members of the flower 
multiply and occupy numerous ranks, the 
receptacle enlarges or lengthens to give them 
insertion or standing-room. 

389. Of elongated forms of receptacle, 
Magnolia and Liriodendron or Tulip-tree give 
familiar instances. The lengthening in the 
former is mainly for the support of both an- 
dreecium and gyncecium ; in the latter, as in 
Myosurus, mainly for the gyncecium only. 
The fall of the matured carpels reveals it 
as a very slender or bodkin-shaped pro- 
longed axis. Of broadened forms, the Straw- 
berry, even in blossom, affords a familiar 
example. (Fig. 406.) In the same order, 
Rubus odoratus shows a very broad and flat 
receptacle: in roses, it is so deeply concave as to become the 
reverse of the strawberry (Fig. 407), being urn-shaped with a 
narrow mouth, upon which the petals and stamens are borne, 


FIG. 405. Stamen of Larrea Mexicana, with a couspicuous ligulate appendage at 
the base within. 

FIG. 406. Receptacle of a strawberry in longitudinal section. 407. Same ofa rose, 
in diagram. 


912 THE FLOWER. 


while the pistils line the walls of the cavity, the base or centre 
of this cavity answering to the apex of the strawberry. 

390. Sometimes internodes are lengthened between certain 
members. In Schizandra, the receptacle, barely oblong in blos- 
som, lengthens greatly in fruiting, so as to scatter the carpels 
on a long filiform axis. 

391. In many Gentians, in Stanleya and Warea among 
Cruciferze, and in most species of Cleome, the internode of the 
receptacle between stamens 
and pistil is developed into 
a long stalk to the latter. 
Gynandropsis (Fig. 409) is 
like its near relative, Cleome, 
except that this very long stalk 
has the lower part of the 
stamens adnate to it: the in- 
ternode between the corolla 
and calyx is broad and slightly 
elevated (or in Cleome, &e., 
narrower and longer) ; and 

He iF so the several floral circles 
are as it were spaced apart by this unusual development of 
receptacular internodes. In Silene (Fig. 408) and many other 
plants of the Pink family, an internode between the calyx and 
corolla is prolonged into a stalk or Stipe." 


1 Srrpe is the general name of a stalk formed by the receptacle or some 
part of it, or by a carpel. To distinguish its particular nature in any case, 
the following terms are more or less employed : — 

THECAPHORE, for a stipe which belongs to a simple pistil itself (where 
it is homologous with a petiole), and is no part of the receptacle, as in Coptis 
or Goldthread. 

GyNopruHore, where the stipe is an internode of receptacle next below the 
gynecium, as the pod-stalk in some Cruciferae, Cleome, and Gynandropsis. 

GonopPuoreE, when it elevates both stamens and pistil, as it seemingly 
does in the lower stipe of Gynandropsis, Fig. 409. 

ANTHOPHORE, when the stipe is a developed internode between the 
calyx and corolla, as in the Pink family, Fig. 408. 

GYNOBASE is a term properly applied to a short and comparatively broad 
portion of receptacle on which the gynecium rests, as in Rue and Orange 
(Fig. 414), Houndstongue, 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 Umbellifere, Fig. 412. 

FIG. 408. Section of a flower of Silene Pennsylvanica, showing the stipe or 
anthophore. 


FIG. 409, Flower of Gynandropsis, with floral circles separated on the elongated 
receptacle. 


FORMS OF THE RECEPTACLE. 213 


392. Instead of forming a stalk, the elongation may 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 Umbelliferee, Fig. 412. In 
Geranium, this prolongation of receptacle 


410 


4i4 


extends far above the ovaries as a beak, to which the styles are 
adnate for most of their length. 

393. In Nelumbium (Fig. 415), 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 perigynous, as in Mae 
Rhamnus (Fig. 415,416) and Cherry (Fig. 357): when carried 
by complete adnation up to the summit of the ovary, it is ep7gy- 
nous, as in Cornus, in Umbelliferee, &c. Not rarely it divides 
into lobes, as in Vitis (Fig. 379, 380), in Periwinkle and most 
Apocynaceous plants, and in Cruciferee. These are termed g/ands 
of the disk, and indeed are commonly glandular or nectariferous. 


FIG. 410. Gyneecium of Geranium maculatum. 411. Thesame 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 Nelumbium, 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. 

395. 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. 
Underthat view, the 
receptacle, instead 
of convex or protu- 
berant, is here con- 
cave, has grown up 
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 first proposed by Link, who intro- 
duced the appropriate name of Hypanrnum. A hypanthium or 
hypanthial receptacle is, as the name betokens, a flower-axis or 
receptacle developed mainly under the calyx. The name is a 
good one, in any case; and such structures as those of Calycan- 
thus (Fig. 417-419), a rose, a pear (the lower part of which is 
evidently an enlargement of peduncle), and of Cactus-flowers 
(Fig. 317), although quite compatible with the theory of adnation, 
are more simply explained by it.? 


1 But, whether the cases are well distinguishable or not, it by 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 

FIG. 417. Flowering branch of Calycanthus. 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. Certrars 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.!. 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.? 


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? Van Tieghem, in his Anatomie Comparée 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 Bau und der 
Befruchtung der Blumen, Berlin, 1793. Even earlier, Kelreuter ( Vorlaufige 
Nachricht, ete., 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 gon 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- 


216 THE FLOWER. 


397. The subject, here considered as a part of morphology, 
must be fully treated, as regards acts and processes, under physi- 
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. 

398. Linnzeus and his successors taught that the adjustments 
in hermaphrodite flowers were such, on the whole, as to secure the 
application of the pollen of its stamens to the stigma of its pistil 
or pistils. The present view is, that this is doubtless strictly 
secured in certain flowers of a moderate number of species, but 
never in all the flowers of any such species; that in ordinary 
flowers, where it may commonly take place, it is not universal ; 
that in the larger number of species there is something or other 
in the floral structure which impedes or prevents it. Some 
flowers are adapted for close fertilization ; some for cross fertili- 
zation; some for either. Here two terms need definition, viz. : 

Close fertilization or Self-fertilization, or Autogamy, the applica- 
tion and action of a flower’s pollen upon its own pistil ; 

Cross fertilization, or Allogamy, the action of the pollen of one 
flower on the pistil of some other flower of the same species. 
This may be near, as when between flowers borne in the same 
cluster or on the same plant; remote, when between flowers of 
distinct plants of the same immediate parentage ; most remote, 
when between different races of the same species. Any thing 
beyond this is hybridization, or crossing of species. 


§ 2. ADAPTATIONS FOR ALLOGAMY OR INTERCROSSING. 


899. The doctrine now maintained appears to have been first 
propounded by Sprengel in the statement that ‘* Nature seems 
to have wished that no flower should be fertilized by* its own 
pollen,” —a proposition which is not wholly tenable, for there 
are blossoms specially adapted to self-fertilization. It was re- 
affirmed in our day by Darwin, in a similar adage, ‘* Nature 
abhors perpetual self-fertilization,’”” — a metaphorical expression 
to which no effective exception has been taken. And the infer- 
ence was drawn by him, that some important good to the species 
must result from propagation through the union of distinct 
individuals, and especially of individuals which have been dis- 
tinct for several or many generations. 


hending Geitonogamy, fertilization by pollen of other flowers of the same 
plant, and Xenogamy, by pollen from a flower on another plant. 


ADAPTATIONS FOR INTERCROSSING. yr} i 


400. The actual proposition, simply stated, is that flowers are 
habitually intercrossed, and that there are manifold structural 
adaptations which secure or favor intercrossing, to such extent 
as to justify the proposition. The proof of the proposition is an 
induction from a very great number of particular observations. 
That intercrossing is beneficial is a rational inference from the 
array of special adaptations for which no other sufficient reason 
appears, or (to resume the metaphor) from the vast pains which 
seem to have been taken to secure this end. ‘This inference has 
been to some extent confirmed by direct experiment.* 

401. Separation of the sexes is a direct adaptation to inter- 
crossing, rendering it necessary between individuals in dicecious, 
and largely fay oring it in most moncecious and polygamous 
flowers. Strictly close fertilization can occur in hermaphrodite 
flowers only ; but it is in these that the most curious adaptations 
for intercrossing are revealed. 

402. The agencies to the one or the other of which most 
flowers are structurally adapted in reference to intercrossing are 
mainly two; viz., the winds and animals, of these chiefly insects. 
Delpino has accordingly classified flowers into Anemophilous and 
Entomophilous ; literally wind-lovers and insect-lovers, but de- 
noting wind-fertilized and insect-fertilized, according to the 
agent by which pollen is transported.* There are hermaphrodite 
and unisexual flowers of both classes, but most wind- ee 
flowers are unisexual. 

403. Wind-fertilizable or anemophilous flowers are mostly neu- 
tral or dull in color, destitute of odor, and not nectariferous. 
Their principal structural adaptations to this end, besides the 
separation of the sexes in most of them, are the superabundance, 
incoherency, dryness, and lightness of the pollen, rendering it 
very transportable by wind and currents of air. The immense 
abundance of pollen, its lightness, and its free and far diffusion 
through the air in Pines, Firs, Taxodium, and other Conifers, 
are familiar. Their pollen fills the air of a forest during anthe- 
sis; and the ‘* showers of sulphur,” popularly so-called, the 
yellow powder which after a transient shower accumulates as 
a scum on the surface of water several or many miles from the 


1 Darwin, The Effects of Cross and Self-Fertilization in the Vegetable 
Kingdom, London, 1876. American Edition, New York, 1877. 

2 Ornithophilous, i.e. bird-fertilized, flowers are to be ranked with entomo- 
philous. The large blossoms of Trumpet Creeper (Tecoma radicans) and 
of Trumpet Honeysuckle (Lonicera sempervirens), and others, are commonly 
visited and probably fertilized by humming-birds as well as by moths; and 
other birds are known to play a similar part in equatorial regions. 


218 THE FLOWER. 


nearest source, testifies to these particulars. All amentaceous 
trees (Willows excepted), Hemp, Hops, &c., are wind-fertilized ; 
and, among perfect flowers, those of most Grasses, Sedges, and 
Plantago. In the latter families especially, the anthers are pro- 
truded or hung out in the air only when just ready to discharge 
their pollen, and are at that moment suspended on suddenly 
lengthened capillary drooping filaments, fluttering in the gentlest 
breeze ; and the stigmas 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- 
able 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-fertilizable or entomophilous 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. Some insects, moreover, visit 
flowers for their pollen, a highly nutritious 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 CSnothera), 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, limbs, 
or bodies of insects, especially to their rough surfaces; and 
in two families (Orchidaceze, Asclepiadacez) 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.! 

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 dicecious flowers, this is necessitated by the separation ; 
in moneecious and polygamous flowers, of various kinds and 


1 Thus nearly every Orchid genus but one has a persistently glutinous. 
stigma ; in the exceptional one, Cypripedium, 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 ora 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.* 

407. Dichogamy, a term introduced by 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 dicecious or mon- 
cecious 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 ; 

Proterogynous (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 
nearly so, is however made to secure the same result through 
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 
slong with it the fact that strictly terminal and also vertical flowers, whether 
erect or suspended, are seldom irregular, while comparatively 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, &e. 

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. 


220 THE FLOWER. 


spikes: a day or two afterwards, the corolla opens, the filaments 
greatly lengthen, and the four 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 case. 
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 


420 


and enclosed. Scrophularia is a good instance of proterogyny 
in flowers fertilized by bees. The flower is irregular (Fig. 
420422), 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. 

409. Proterandry. The process is the reverse, and is at- 
tended with much more extended movements in Clerodendron 
Thompsoniz, 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. aA | 


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 strongly to one side, out of the way 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’ observed that the anthers of freshly 


1 In Meehan’s Gardeners’ Monthly, September, 1878, 278. 


FIG, 423. Flower of Clerodendron Thompsoniz, 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, Epilobium angustifolium or E. 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 


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 very 
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 
j anthers of a Cam- 
panula discharge all 
their pollen in the 
unopened bud, and it 
is nearly all deposited 
on the style which 
they surround, the 
upper part of which 
is clothed with a coat 
of hairs for holding 
= ies the pollen. (Fig. 427.) 

In the open flower, the stamens are found to be empty and withered, 
as in Fig. 428. These flowers are visited by bees and other insects 
for the 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 Epilobium angustifolium or spicatum; in the first, 
freshly expanded; in the second, a few days older. 

FIG. 427. 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 
style. (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 Composit, 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 the natural size. 


pct 8 THE FLOWER. 


margin (ray-flowers, with ligulate 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- 
alescent anthers. The pollen is early discharged into the interior 
of this tube. The style, with somewhat enlarged and brush-like 
tip, at first 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 


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. 435), exposing to other 
pollen the stigmatic receptive surface which until now was un- 
approachable. 

412. In Parnassia, which has sessile stigmas, their receptive 
surface is actually not formed until the anthers become effete ; 


FIG. 432. A ligulate female flower of the same, and a central hermaphrodite flower. 
433. Upper part of the latter, more enlarged, the tube of anthers projecting from the 
corolla, and the pollen projecting from apex of the anther-tube, being pushed up by the 
lengthening of the style beneath. 434. This style now projecting, and some pollen still 
resting on its tip. 435. Tip of same style (more advanced and magnified); the two 
branches spreading, still carrying some pollen on the apex of each arm or branch, and 
by the “ivergence now exposing the stigmatic inner faces. 


ADAPTATIONS FOR INTERCROSSING. 225 


and, as the plants or stems are single-flowered, they are function- 
ally dicecious while structurally hermaphrodite. 

413. The adaptations for hermaphrodite intercrossing with 
synanthesis (407), 7. e. where there is no essential difference of 
time in the maturing of anthers and stigma, are manifold. 
They may be classed into those without and those with dimor- 
phism of stamens and pistils, or, in other words, those with 
Homogonous and those with Heterogonous flowers.} 

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, 358) — 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 Amer. Jour. Sci. ser. 5, xiii. 82, and in Amer. 
Naturalist, January, 1877. Dimorphism in flowers may affect the perianth 
only, and not the yovj or essential organs; or there may be two kinds of 
flowers as respects these also, but with no reciprocal relations, as in cle/sto- 
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 authers and stigma 
thus brought up against the bee’s abdomen. 439. Style and stigma, with part of the 
ovary, more magnified, a fringe of fine bristles around the stigma. 


15 


226 THE FLOWER. 


Wistaria (Fig. 456-439), in which the light fringe of stiff hairs 
around the stigma (shown in Fig. 439) would not prevent pollen 
of surrounding anthers from falling upon it. Yet when a bee 
alights upon the keel, 
with head toward the 
base of the flower, and 
proboscis is inserted 
for nectar between the 
foot of the standard 
and the keel, the latter 
is depressed by 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 

443 bee. This does not bring out the anthers as in 
Wistaria, but these remain untileffete 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 flower, which 
flower immediately gives to the same spot some of its pollen, to 
be transferred to the next flower’s stigma, and so on. 

417. Two special modifications of the papilionaceous type 


FIG. 440. Flower of Robinia hispida, the standard and wings removed. 441. Same, 
as depressed by the weight of a bee, causing the stigma and pollen-laden tip of the style 
to protrude. 442. Enlarged section of same in the bud, leaving one keel-petal, half the 
stamens, and the pistil in view. 443. Style and stigma at a later period, the beard 
loaded with pollen; more magnified, 


ADAPTATIONS FOR INTERCROSSING. 227 


need particular mention. One of them, the Bean-blossom, is 
weil known to botanists; the other not so. The peculiarity in 
the common Bean, 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 early discharge their pollen, much of which adheres 


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 remoyed. 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 
self-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 
are 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 
behind, or a shallow sae as seen 

ees 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 


452 453 
standard. (Fig. 450, 452,453.) So it remains if untouched until 
the blossom withers: no self-fertilization has ever been observed, 
and none ordinarily occurs. 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. 


FIG 450. Flower of Apios tuberosa, unvisited. 451. Same after visitation, the 
keel dislodged from the retaining notch, and more incurved; the tip of the style pro- 
truded and thrust forward, followed by the anthers. 

FIG. 452. Enlarged vertical section of flower-bud of Apios tuberosa. 453 A flower 
with half the standard cut away, to show the blunt apex of the keel resting in the 
notch. 454. Diagram of flower, with half of the standard cut away, toshow what takes 
place when the apex of the keel is liberated. The figures (also those from 423 to the 
present), and the first 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 honey-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 body, 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. 


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 
(externally 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 — 4s 

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. 


250 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 flower to flower, 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 style 
(Fig. 459): 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 faces 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 Asclepiadaceze and the Orchidacee. 
While in the Iris family the number of stamens is reduced from 
six to three, in all the Orchis family, except Cypripedium, the 
stamens are 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 face 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 bee’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 petaloid style- 
lobe and stigma cut away. The section of the stigma is seen edgewise: the rough 
upper surface only is stigmatic. 


ADAPTATIONS FOR INTERCROSSING. 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 isin Orchis 
and in the commoner re- 
presentatives of Orchis in 
North America (viz. Ha- 
benaria, &c.) that the 
most exquisite adapta- 
tions are found, and the 
greatest economy — se- 
cured ; paralleled, how- 
ever, by most of the very 
humerous and various epi- 
phytic and by various ter- 
restrial Orchids of warmer 
regions. <A single illus- 
tration may here suffice ; 
and Darwin’s volume on 
the Fertilization of Or- 
chids (596, 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 Habenaria 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 nectary), in which nectar is 


1 


1 
Y 


FIG. 460. Flower of Habenaria or Platanthera orbiculata, enlarged. 461. Combined 
stamen and stigma, more enlarged. 462. One of the two pollen-masses (pollinia), with 
its stalk and glutinous disk or gland. 462¢. Lower part of this stalk and its disk, more 
magnified. 


232 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 below: most of the 
lower part of the space between is excessively 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 
B elastic tissue, by which they are 

all attached to a firmer central 
stalk, or caudicle. (Fig. 465-465.) 
To the lower end of this caudicle 
(directly to the end of it in our 
Habenariz and Orchises gener- 
ally, in this instance to the inner 
side of the 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. 461. The exposed 
face of the disk being covered with 
463 465 a durable layer of very viscid mat- 

ter, the body itself is sometimes termed a gland, and not improperly. 
The viscidity is nearly of the same nature as that of the interven- 
ing stigma, of which the glands are generally supposed to be 
detached portions. If so, then a portion of the stigma is cut off 
from the rest and specialized to the purpose of conveyance of the 
pollen. When a finger’s end or any smaller body is touched to 
these disks, they adhere so firmly that the attached pollinia or 
pollen-masses are dragged out of the cell and carried away en- 
tire. Some of these pollen-masses have been found attached by 
the disk to the eyes of a large moth. When a moth of the size 
of head and length of proboscis of Sphynx drupiferarum visits a 
spike of these flowers, and presses its head into the centre of the 


FIG. 463. A more magnified pollen-mass of Platanthera orbiculata, with its stalk 
and gland. 464. Five of the separate portions or pollen-packets, with some of the 
elastic threads of tissue connecting them. 465. A portion more highly magnified, with 
some of the pollen-grains in fours detached. 


ADAPTATIONS FOR INTERCROSSING. 233 


flower so that its proboscis may reach and drain the bottom of 
the nectariferous tube, a pollen-mass will usually be affixed to 
each eye: on withdrawal, 
these will stand as in Fig. 
466. Within a minute they 
will be turned downward 
(Fig. 4667), not by their 
weight, but by a contraction 
in drying of one side of the 
thick piece which connects 
the disk with the stalk. 
When a moth in this con- 
dition passes from the last 
open flower of one spike to 
that of another plant, and 
thrusts its proboscis down a 
nectary, the transported pol- 
len-masses will be brought 
in contact with the large 
glutinous stigma: on with- 
drawal, either some of the 
small pellets of pollen will be 
left adherent to the stigma, 
the connecting elastic threads 
giving way; or else a whole 
pollen-mass will be so left, 
its adhesion to the glutinous aoe 
stigma being greater than that of the disk to the moth’s eye. 
The former is a common and a more economical proceeding, as 
then a succession of flowers are abundantly fertilized by one 
or two pollen-masses. In either case, new pollen-masses are 
earried off from fresh flowers and applied to the fertilization 
of other blossoms on the same and eventually on those of differ- 
ent individuals. Cases like this, and hundreds more, all equally 
remarkable, serve to show how sedulous. sure. and economical are 
the adaptations and processes of Nature for the intercrossing of 
hermaphrodite flowers. 

422¢, An arrangement analogous to that of Orchids, and 
similarly subservient to cross-fertilization, characterizes the 
otherwise widely unlike Asclepias family. In Asclepias (Milk- 
weed) there are five stamens surrounding a large stigmatic 


FIG. 466. Front part of Sphynx drupiferarum, bearing a pollen-mass of Platan- 
thera orbiculata affixed to each eye, in the early position. 466. Front view of the head, 
later, showing the pollen-masses deflexed. 


oa 


Zot THE FLOWER. 


body, and alternating with these five two-cleft glands, the ver- 
tical chink or groove of which is glucinous. To each gland is 
firmly attached, by a caudicle or stalk, a pollen-mass of an ad- 
jacent anther. (Fig. 522.) <A slight force upraising the gland 
detaches it from the stigma and drags the pair of suspended 
pollen-masses out of their cells. Insects visiting the blossoms 
commonly dislodge them, the gland adhering to their legs or 
tongues when these happen to be drawn through the adhesive 
chink, and convey them from one flower to another. Without 
such aid the flowers of Asclepias ravely set seed.' 

423. Dimorphism, 7. e. the case of two kinds of blossoms, both 
hermaphrodite, on the same species, is another adaptation to 
intercrossing. Not all dimorphism, however, for in eleistogamous 
dimorphism (484) the intent to self-fertilize is evident. There 
may also be dimorphism as to the perianth, not particularly 
affecting fertilization. One kind, however, and the commonest, 
is a special adaptation to intercrossing, viz. : 

424. Heterogonous Dimorphism. (415, note.) This term is 
applied to the case in which a species produces two kinds of 
hermaphrodite flowers, occupying different individuals, the flowers 
essentially similar except in the andreecium and gyneecium, but 
these reciprocally different in length or height, and the adapta- 
tions such that, by the agency of insects, the pollen from the 
stamens of the one sort reciprocally fertilizes the stigma of the 
other.?, This dimorphism has been detected in about forty genera 
belonging to fourteen or fifteen natural orders, widely scattered 
through the vegetable kingdom ; but there are far more examples 
among the Rubiacez than in any other order. Sometimes all 
the species of a genus are heterogonous, as in Houstonia, and 


1 The reported sensitiveness of the gland, referred to in the first issue of 
this volume (1879), was founded upon misinterpreted observations. 

2 This peculiar arrangement has been long known in a few plants, such 
as Primula veris, P. grandiflora, and Houstonia. In Torrey and Gray’s 
Flora of North America, ii. 38, 39 (1843), these flowers are said to be dicecio- 
dimorphous, not’ denoting that they are at all unisexual, but 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 Dimorphic 
Condition in the Species of Primula, and on their. Remarkable Sexual Rela- 
tions,” published in the Journal of the Linnean Society, vi. (1862), 77: repub- 
lished, in 1877, as the leading chapter of his volume entitled “The Different 
Forms of Flowers on Plants of the Same Species.” Mr. Darwin had termed 
these flowers simply Dimorphic; but in this volume he adopted Hilde- 
brand’s name of Heterostyled for this kind of blossom. The difference, 
however, affects the andreecium, and even the pollen, as well as the style; 
wherefore we proposed for it the name of Heteroyonous or Hetereyone dimor- 
phism, as mentioned in a former note, 413. 


ADAPTATIONS FOR INTERCROSSING. 235 


Cinchona, sometimes only a part of them, as in Primula and 
Linum. In Hottonia, a Primulaceous genus of two species, the 
European one has heterogonous dimorphism’ for cross-fertiliza- 
tion: the American one has homogonous showy flowers with 
only the general chance for intercrossing, and earlier flowers 
which are cleistogamous for self-fertilization. 

425. The nature of heterogone dimorphism may be well under- 
stood from a single example. The most familiar one is that of 
Houstonia ; but, in larger blossoms, Gelsemium is a fine illus- 
tration in the Southern United States, and Mitchella (Fig. 467) 
mostly in the Northern. Raised from the seed, the individuals 


are about equally divided between the two forms: namely, one 
form with long style and short or low-inserted stamens; the 
other with short style and long or high-inserted stamens. The 
stigmas in one rise to about the same height as the stamens in 
the other, both in the tall or exserted organs and in their low 
and included counterparts, as is shown in Fig. 468, answering 
to the left hand and Fig. 469 to tle right hand flowers of Fig. 467. 
A bee or other insect with proboscis of about the length of the 
corolla-tube, visiting the blossoms of Mitchella, will brush the 
same part of its body against the high anthers of the long- 
stamened and the high stigmas of the long-styled forms; and 


> -  r-O 


1 ©. C. Sprengel, as Darwin mentions, had noticed this, before 1793. He, 
“with his usual sagacity, adds that he does not believe the existence of 
the two forms to be accidental, though he cannot explain their purpose.” 
Darwin, Forms of Flowers, 51. 

Some heterogonous Primulas are said to produce homogonous varieties in 
cultivation. In Primula, and in other genera, there are species which seem 
as if of one sort only, no reciprocal sort being known, as if one form had 
become self-fertile and the other had disappeared. 


FIG. 467. Partridge Berry, Mitchella repens, in the two forms, viz. long-stamened 
and short-styled, and short-stamened and long-style. : 


236 THE FLOWER. 


the same part of the proboscis against the low anthers of the 
short-stamened and the low stigmas of the short-styled form. 

426. Moreover, Dar- 
{i win has ascertained by 
microscopical examina- 
tion that the pollen of 
the two differs in size or 
shape, and by experi- 
ment that it is less 
active upon its own 
stigma than upon the 
other; indeed, that in 
many cases (as in some 
species of Linum) it is 
quite inactive or impo- 
tent not only upon its 
own stigma but upon its 
own-form stigma, while 
it is prepotent on the other, and this reciprocally of the two 
forms.’ Here, then, are flowers structurally hermaphrodite, but 
functionally as if dicecious, securing all the advantages of the 
latter, along with the economical advantage that both sorts of 
individual and every blossom may bear seed. With dicecism 
only about half the plants could be fruitful. 

427. Heterogonous Trimorphism. A threefold heterogonism 
is known in certain species of a few genera; and this complica- 
tion may have certain conceivable advantages over dimorphism. 
Where seedling dimorphous individuals are few and far between 
(those multiplying from root would all be alike), there would 
be an even chance that any two near each other were of the 
same form and therefore sterile or imperfectly fertile. But if 
the organization were 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 different 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 


Ss 


QS = 


1 Impotence of own pollen, either absolute or relative. occurs no less in 
certain flowers which are not dimorphous, as in Corydalis, some species of 
Passiflora, &c. On the contrary, many dimorphous flowers are in a certain 
degree self-fertile, especially in the long-stamened and short-styled form. 
These subjects are physiological, and belong to another volume. 


FIG. 468. Long-styled flower of Fig. 467, laid open. 469. Long-stamened flower of 
the same laid open. Both equally enlarged. 


ADAPTATIONS FOR INTERCROSSING. 237 


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 pistilin 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 .vica. 
organs, all as distinct 
from one another as if 
they belonged to different 
species; and, if smaller. 
functional differences are = *!e4 
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 Tn an article On the Sexual Relations of the Three Forms of Lythrum 
Salicaria, in Jour. Linn. Soe. 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.) 


a 


238 THE FLOWER. 


false coadaptation.” The whole arrangement is displayed in 
the annexed diagram (Fig. 470), and in the following account 
of the operation.! ‘* Ina state of nature, the flowers are inces- 
santly visited for their nectar by hive and other bees, various 
Diptera, and Lepidoptera. The nectar is secreted all round the 
base of the ovarium; but a passage is formed along the upper 
and inner side of the flower 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 brushed by 
the lower hairy surfaces of the insects’ bodies. The shortest 
stamens, which lie 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, and 
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 I 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 flower allows the easiest access to it, 
as in Lythrum. . . . 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 flowers, the anthers of the longest stamens, 
bearing the green pollen, are rubbed against the 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 flower. 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, &. 


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 days 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 polien, 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 ‘ legitimate 
union,” z.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. Nessa 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,’ with results 
quite as decisive as in Lythrum Salicaria. One genus of 
Monocotyledons has trimorphous blossoms, viz. Pontederia, of 
which the North American P. cordata is a good illustration.’ 

431. All known flowers exhibiting reciprocal dimorphism or 
trimorphism are entomophilous: no such wind-fertilized species 
is known. Few of them are irregular, and none very irregular : 
they do not occur, for instance, in Leguminosz, Labiate, 


1 Monatsber. Akad. Berlin, 1866; Bot. Zeit. 1871, &ce. 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, tliree in each set. 


240 _THE FLOWER. 


Scrophulariaceze, Orchidacerw, &c. Nature is not prodigal, and 
does not endow with needless adaptations flowers which are 
otherwise provided for. 


§ 3. ADAPTATIONS FOR CLOSE FERTILIZATION. 


432. Even where cross-fertilization in bisexual flowers is 
obviously arranged for, 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 be 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 proterandrous, 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, they remain for some days 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- 
fertilize even in the bud in early spring, when insects are scarce, 
but are habitually intercrossed by insectsin summer. Somewhat 
similarly, according to Hermann Mueller.’ certain species, such 
as Euphrasia officinalis and Rhinanthus Crista-galli, habitually 
produce two kinds of blossoms, one larger and more showy, 
usually affecting 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 der Blumen durch Insekten, 204; 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 Linnzeus,! 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 Malpighiaceze 
in 1832, and recorded in 18435 that Adolphe Brongniart had well 
investigated those of Specularia, and that Weddel had discoy- 
ered them in Impatiens Nolitangere. <A full account of the then 
known cases was given by Mohl? in 1863; but D. Mueller, of 
Upsala, who examined Viola canina, is said by Darwin to have 
given,” in 1857, *: the first full and satisfactory account of any 
cleistogamic flower.” The appropriate name of cleistogamous 
was given by Kuhn,’ 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 Dicotyiedons. All but the 
Grasses*® 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. Linneus 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 cleistogenous, which is not so proper. We prefer cleistoyamous to 
cleistogamic (and so of similar terms), as best harmonizing with the Latin 
adjective form, both in form of termination and in euphoniously taking the 
accent upon the antepenult. 

® 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 


» 


242 THE FLOWER. 


increase is by cleistogamous self-fertilization, which thus offsets 
the incidental disadvantage of the former mode. 

437. In general, the cleistogamous are like unto the ordinary 
flowers arrested in development, some arrested in the almost 
fully formed bud, 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 being 
rudimentary, they are singularly inconspicuous. Consequently, 
insects do not visit them; nor, if they did, could they find an 
entrance. Such flowers are therefore invariably self-fertilized ; 
yet 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.”! 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 always 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 
intererossing, — one should compare the small number of pollen- 
grains which so completely serve the purpose in a typical cleis- 
togamous flower (say 400 in Oxalis Acetosella, 250 in Impatiens, 
100 in some Violets) with the several thousands of all entomo- 
philous cross-fertilized flowers, rising to over three and a half 
millions in the flower of a Peony, also their still greater number 
in many anemophilous blossoms. To this loss should be added 
the cost of a 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 CAaLyx 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 (zstivation), have already been considered in Chap. IV. 
Sect. I., Il. And most of the morphology of calyx 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 Labiatee, 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, Helianthemum, 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 equally 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 Cactacee. 

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) ; Zriphyllous, of 
three ; Tetraphyllous, of four ; Pentaphyllous, of five ; Hexaphyllous, 
of six, and so on. A tulip and a Tradescantia flower have a 
hexaphyllous perianth, but composed of two circles, answering 
to calyx and corolla; each Triphyllous.? When the character 


1 Perianthium, alias Perigone or Perigonium. (296.) 

2 As elsewhere explained, when numerical composition is indicated without 
reference to nature of parts, the terms d/merous, trimerous, tetramerous, penta- 
merous, &c., may be used. (522.) 


244 THE FLOWER. 
of the organ, 7. e. whether calyx or corolla is to be specified, 
the word sepal or petal is employed in the combination ; as, 

Disepalous, of two sepals ; T'risepalous, of three; Tetrasepalous, 
of four; Pentasepalous, of five (also written 5-sepalous, and ac- 
cordingly 2-sepalous, 3-sepalous), and so on: also, 

Dipetalous, Tripetalous, Tvtrapetalous, Pentapetalous (2-5- 
petalous), &c., when the corolla is concerned. 

442, Monophyllous, Monosepalous, and Monopetalous are the 
proper terms for perianth (calyx, corolla, &c.) composed of a 
single leaf. Likewise Polyphyllous, Polysepalous, and Polypetal- 
ous for the case of a considerable but unspecified number of 
members. Unfortunately, in the Linnean and long-prevalent 
use, monopetalous was the term employed to designate a corolla 
of one piece in the sense, or the fact, of a coalescence or grow- 
ing together of two, three, five, or more petals into a cup or 
tube ; and so of a calyx, of a whorl of bracts, &e. And poly- 
petalous, polysepalous, and polyphyllous were the counterparts 
of this, meaning of more than one distinct piece, whatever the 
number. The misleading use, consecrated by long prescription, 
is not yet abandoned, but will in time be obsolete. In present 
descriptive botany, a polyphyllous calyx, or a_ polypetalous 
corolla, or a 5-petalous 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. (329.) 

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, Gamosepalous for the calyx ; 
these terms meaning united petals or sepals. The older and mis- 
leading names JMonopetalous and Monosepalous, although current 
up to a recent day, should be discontinued. Another term is 
not rarely used in Germany, that of Sympetalous, for the gamo- 
petalous (or formerly monopetalous) corolla, — therefore Syn- 
sepalous for a similar calyx. It is perhaps a more apt term 
than gamopetalous, 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 Po/ypetalous, as 
already explained. (442.) Itis adopted by Eichler, &e.  Chori- 
sepalous is the term applied to the calyx. Déalypetalous (em- 
ployed by Endlicher) 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. Hleutheropetalous (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 
(3-parted, 5-parted, &c.), when the sinuses extend almost to 
the base ; cleft, when about to the middle ; /obed, a general term 
for any considerable separation beyond toothing ; dentate or 
toothed (3-toothed, 5-toothed, &c.), when the union extends 
almost to the summit; entive, 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 Lamrya or BLape: any much contracted 
base is the Uncuts or Ctaw. The latter is very short in a rose- 


471 472 473 


petal, but long and conspicuous in a pink and all flowers of that 
tribe (Fig. 471), in many Capparidez (Fig. 409) and Crucifere. 
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 TuBr: an expanded terminal portion, either 
divided or undivided, is the Lime or Borper. The limb may 


FIG. 471. Corolla of Soapwort, of five separate long-clawed or unguiculate 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. Flower of the Cypress-Vine (Ipomca Quamoclit); the petals a little farther 
united into a five-lobed spreading border. 

FIG. 474. Flower of the Ipomcea coccinea; the five component petals perfectly united 
into a trumpet-shaped tube, and beyond into an almost entire spreading border. 


246 THE FLOWER. 


be parted (that is, the component parts not united) quite or 
nearly down to the tube or base, as in Fig. 472, 475 ; or less so, 
as in Fig. 473, 476 (with limb 
d-lobed) ; or with merely angles 
or points to represent the tips 
of the component members, as in 
Fig. 474; or with even and entire 
border, as in common Morning- 
Glory, Fig. 482. 

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 Turoat, in Latin Faux, pl. 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 Crown (385, shown 
in Fig. 403, 404, 471) usually belong to the throat of a gamo- 
petalous corolla or perianth, as in Oleander, Comfrey, Borrage, 
Narcissus, &¢., or to a corresponding position when the parts 
are not coalescent. 

449. Forms of Corolla, Calyx, &e.- 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 Leguminose (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 

Caryophyllaccous, or Fink-flower (Fig. 471), a regular corolla, 
of five long-clawed (wnguiculate) petals, the claws enclosed in a 
tubular calyx and the blades spreading ; and the 

Cruciferous, of four somewhat similar petals, the four abruptly 
spreading blades in the form of a cross (erucrate), as in Fig. 394. 

Rosaceous, with roundish and widely spreading petals on very 
short or hardly any claws, as in Rose and Apple-blossoms. 

Liliaceous, a 6-phyllous perianth of campanulate or funnelform 
shape ; the members either distinct, asin most common lilies and 
tulips, or gamophyllous, as in Lily of the Valley. All but the 
first and last of these sorts are examples of regular and chori- 
petalous perianth. 

Orchidaceous flowers are of a peculiar irregularity, combining 
both calyx and corolla: one member, the petal in front of the 


476 


FIG. 475. Rotate or wheel-shaped and five-parted corolla of the Bittersweet (So- 
lanum Duleamara). 


FIG. 476. Whiecl-sheped and five-lobed corolla of the common Potato. 


PERIANTH, OR FLOWER-LEAVES. 247 


stamen and stigma, differs from the rest in shape and in being 
nectariferous (as in Fig. 460) ; it is named the Lapertum. 

Galeate is a term applied to a corolla the upper petal or part 
of which is arched into the shape of a casque or helmet, called 
the Galea; as in Aconite (Fig. 357) and Lamium, Fig. 479. 
In the former the galea is of a single petal; in the latter, it 
consists of two, completely united. 

450. Gamophyllous forms with special names are chiefly the 
following. Illustrations are usually taken from the corolla, but 
the forms and terms are not peculiar to it, excepting the first, 
viz. the 

Ligulate or Strap-shaped corolla (Fig. 288, &c.), which is 
nearly confined to Compositz. Here a corolla, formed of three 
or five petals, imitates a single petal, except at its very base, 
which is commonly tubular: the remainder is as though the tube 
had been split down on the upper side and flattened out. The 
corolla of Lobelia, type of a family most nearly related to Com- 
posite, illustrates this. (Fig. 488.) 


4i7 478 479 480 481 
451. The names of the general forms are mostly taken froim 
some resemblance to common objects. All those in common 
use will be found in the Glossary: a few leading ones are here 
specified. They may be divided into the regular and the irregu- 

lar. The principal irregular form with a special name is the 
Labiate, or lipped, also termed Bilabiate, as there are two lips, 
an upper and a lower (superior and inferior, or anterior and 
posterior, 290), although one of them is sometimes obscure or 
abortive. This bilabiate character in the corolla, and often in 
the calyx also, pervades several orders with gamopetalous 
flowers, and gives name to one of them, the Labiatz, to which 


FIG. 477. Campanulate corolla of the Harebell, Campanula rotundifolia. 478. 
Salverform (hypocraterimorphous) corolla of Phlox. 479. Labiate (ringent) corolla of 
fLamium; a side view 480. Personate corolla of Antirrhinum or Snapdragon. 481. 
Personate corolla of Linaria, spurred (calearate) at the base. 


248 THE FLOWER. 


the Sage and Mint belong. Such flowers are 5-merous, and 
have two members specially united to form one lip, and three in 
the other. The odd sepal being posterior (or next the axis of 
inflorescence) , and consequently the odd petal anterior, the calyx 
has its lower lip of two sepals and its upper of three; while the 
corolla has its upper lip of two petals and its lower of three. 
But in Leguminose, where the calyx is sometimes bilabiate, and 
where the odd sepal is anterior (or toward the bract), this is 
reversed, and two sepals or lobes of the calyx form the upper 
lip and three the lower. A bilabiate corolla is 

Ringent, that is gaping or open-mouthed, when the throat is 
freely open, as in Lamium, Fig. 479 ; 

Personate, or masked, when the throat is closed, more or less, 
by a projection of the lower lip called the PaLatr, as in Antir- 
rhinum and Linaria, Fig. 480, 481. 

452. Of regular forms, there are the following, beginning with 
that having least tube: 

Rotate, or Wheel-shaped (Fig. 475, 476), widely spreading from 
the very base, or from a short and inconspicuous tube. 

Crateriform, or Saucer-shaped, like rotate except that the broad 
limb is cupped by some upturning toward the margin. 

Hypocrateriform, or rather (not to mix Latin and Greek) 
Hypocraterimorphous, in English Salverform, when a rotate or 
saucer-shaped limb is raised on a 
slender tube which does not much 
enlarge upward ; that is, where a long 
and narrow tube abruptly expands 
into a flat or flattish limb, as in 
Fig. 478. In Fig. 472-474 are seen 
salverform corollas with somewhat 
more upwardly dilated  (trumpet- 
shaped) tube. The salver or hypv- 
eraterium, which the name refers to, 
with a stem or handle beneath, is now 
to be met with only in old pictures. 

Tubular, when strictly used, denotes 
a gamophyllous perianth with limb 
inconspicuous in proportion to the 
tube, as in Trumpet Honeysuckle, or as Fig. 472-474 would be 
if the limb were much diminished or wanting. But it is some- 
times used in the sense of having a conspicuous tube. 


FIG. 482. Calyx and funnelform (infundibuliform) corolla of a common Morning- 
Glory, Ipomcea purpurea. 


ANDRGCIUM, OR STAMENS. 249 


Infundibuliform, or Funnelform, such as the corolla of common 
Morning-Glory (Fig. 482), denotes a tube gradually enlarging 
upward from a narrow base into an expanding border or limb. 

Campanulate, or Lell-shaped (Fig. 477), denotes a tube of length 
not more than twice the breadth, moderately expanded almost 
from the base, the sides above little divergent. 


Section VI. THE ANDRG@CIUM, OR STAMENS IN PARTICULAR. 


453. The whole Stamen. For the general character and some 
of the modifications of the stamens, see the first (501) and por- 
tions of the succeeding sections of the present chapter. The 
terms peculiar to these organs, and of common use in botanical 
description, were nearly all coined by Linnzeus, and employed as 
the names of classes in his sexual system. (672.) The sub- 
stantive names of those classes which are characterized by the 
number of stamens, and which were designated by Greek nume- 
rals prefixed to andria (the Greek word for man being used 
metaphorically for stamen), are put into adjective form, as 
follows : 

Monandrous, for a flower with a solitary stamen ; Diandrous, 
for a flower with two stamens; Z7riandrous, with three; Tetran- 
drous, with four; Pentandrous, with five ; Hexandrous, with six ; 
Heptandrous, with seven; Octandrous, with eight ; Lnneandrous, 
with nine; Decandrous, with ten; Dodecandrous, with twelve; 


Polyandrous, with a greater or indefinite number, or /eosandrous 
(meaning twenty-stamened) when a polyandrous flower has the 
stamens inserted on the calyx, as in the Cherry (Fig. 337), 
Pear, &c. 


FIG. 483. Diadelphous stamens (9 and 1) of a Pea. 484. Monadelphous stamens of 
a Lupine. 485. Monadelphous stamens, &c., of Mallow. 
FIG. 486. Five syngenesious stamens of a Composita. 487. The same, laid open. 


250 THE FLOWER. 


Didynamous is a term applied to an andreecium of four sta- 
mens in two pairs, a longer and a shorter, as in Fig. 361. 

Tetradynamous is similarly applied to that of six stamens, two of 
them shorter, in the manner characteristic of Cruciferz, Fig. 396. 

454. Terms which denote coalescence of stamens, whether by 
their filaments or their anthers, are 

Monadelphous, that is, in one brotherhood, by coalescence of 
the filaments into a tube, as in the Mallow (Fig. 485), Lupine 
(Fig. 484), Lobelia (Fig. 488), &e. 

Diadelphous, in two brotherhoods, by coalescence of the fila- 
ments into two sets; sometimes an equal number in each, as in 
Fumariacee (Fig. 390), sometimes nine in one set and one 
separate, as in the Pea (Fig. 483) and most Papilionaceze. 

Triade/phous, with filaments united in three sets or clusters, as 
in Hypericum. 

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 


489 491 492 


(Fig. 486, where they are five in number and the filaments dis- 
tinct), in Cucurbita (Fig. 489, 490, where they are three in 
number and the filaments partly monadelphous), and in Lobelia 
(Fig. 488, where they are also five and the long filaments are 
mainly monadelphous). 


FIG. 488. Flower of Lobelia cardinalis, with tube of corolla divided on one side; 
filaments and anthers united into a tube: f. tube of filaments; a. of anthers. 

FIG. 489. Male flower of Cucurbita (Squash), with limb of calyx and coroHa cut 
away, to show the stamens, viz., three filaments, separate at base but monadelphous 
above, and three syngenesious anthers ina kind of head. 490. Stamens of the same, 
enlarged and the upper part cut away, to show the union. The anthers are sinuous. 
491. A detached stamen of the Melon, with loosely sinuous anther. ; 

Fig. 492. Stamens and style of a Cypripedium, united into one body or co/umn: 
@, anthers; st. enlarged sterile stamen; stig. the stigma. 


ANDRG&CIUM, OR STAMENS. 251 

455. Of terms relating to adnation of stamens, besides the 
general ones of hypogynous, perigynous, epigynous (332), and 
epipetalous, or adnate with corolla, there is the special one of 

Gynandrous, having stamens borne upon the pistil, as in 
Orchidacex. In Cypripedium, 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 consolidated 
into one mass, and there is no evident style. 

456. A complete stamen consists of Fmamenr 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 by its bearing an anther; as in the 
transition states between the true petals and stamens of Nym- 
phzea, 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 theese, 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 Connectrvum, 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 faces forward or backward. In the vast genus Solanum, 


i) 


495 44. 495 45 497 * 498 

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, Blueberry, &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 Lauracez, 

~™ 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. 

461. The attachment of the anther to the filament presents 
three principal modes, which are connected by gradations. 
These are the 

Innate (Fig. 495, 496), 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. 


FIG. 493. A stamen, with its anther, 6, surmounting the filament, a, and opening 
in the normal manner down the whole length of the outer side of each cell. 

FIG, 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. 

FIG. 496. A stamen of Isopyrum biternatum, with innate anther. 497. Stamen of 
Liriodendron, or Tulip-tree, with adnate extrorse anther. 498. Stamen of CEnothera 
glauca, with the anther fixed hy its middle and versatile 

FIG. 499. A stamen of Asarum Canadense, with adnate anther and prolonged tip 
to connective. 


ANDRGECIUM, 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, Limodendron 
(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 anthesis 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. tarned inward, or Anticous, when it faces toward 
the axis of the flower; as in Nymphzeacee (Fig. 318), in Violet 
and Lobelia (which are adnate and introrse), and in Génothere. 
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 
always 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 bas/fixed or med/ fired, 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. 


954 THE FLOWER. 


465. The connective may be appendaged either by a prolon- 
gation or otherwise from the tip (as in Fig. 499), or from the 
back, as in Violets and in many Ericaceous plants. 

466. The normal anther is two-celled, bilocular, or (to use 
a less common term) didiecous, and its lobes or cells parallel, 
right and left; but the cells at first, and sometimes at maturity, 
are bilocellate, that is 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 
B | in many others, this 
Y \ine of dehiscence is 

marginal or lateral, 

either  stricily or 
a ne =e nearly so, as in 
Fig. 500. When introrse or extrorse (as in Fig. 501, 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 
the two locelli on the same side of the midrib or connective 
(right and left) are usually confluent into one pollen-filled cavity 
or cell at maturity if not earlier, or at least the partition between 
them 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 quadrilocellate, 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 1s homologous with the blade or 
an apical portion of the blade, therefore the two lobes or thecz 
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 This is the view long ago taken by Cassini and Reeper, and it may still 
be maintained as the best morphological conception. Mohl interposed some 
objections to its universality ; but, as presented in Sachs’s Text-Book, they 
are not incompatible with the common morphology. Sachs takes the fila- 
ment with the connective to be the homologue of the whole leaf, and the 
anther-cells as appendages. Others, in likening the anthers to glands, adopt 
a similar view. 


FIG. 500. Innate anther, same as Fig. 496. in younger state, with transverse section, 
showing the four locelli. 501. Same of an adnate extrorse anther, such as Fig. 497. 
502. Same as the preceding but mature and dehiscent, the two locelli becoming one cell 
hy the vanishing or breaking up of the partition. 


ANDRGECIUM, 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 many 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 may, in their carly 
development, become confluent at the apex, as in the 5s 
Mallow family (Fig. 506), 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 by confluence. 
In another way, the anthers of some species of Orthocarpus (gen- 
erally resembling Fig. 505, but the lobes or cells quite distinct 
or even separated at apex) lose one of the cells by partial or 
complete non-development and so become one-celled by abor- 
tion. The anther of Gomphrena (Fig. 507) is completely uni- 
locular by abortion or suppression of the companion cell. Thus 
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, asin 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 


oH 50D 506 HOT 


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] art of the anther, 
in section, the upper a part of a leaf. 

FIG. 504. Stamen of Menispermum Canadense, the quadrilocellate anther divided. 

FIG. 505. Stamen of Pentstemon pubescens, 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 of Gomphrena or Globe Amaranth, medifixed, of a single cell, 
dehiscent. 


256 THE FLOWER. 


proper filament, and bearing an anther-cell at each end. Ina 
few species, the two anther-cells are nearly alike; in more, the 
lower one is imperfect, as in Fig. 510%; in more, it is abortive 
or wanting altogether, as in Fig. 510°. Then, in the related 
Californian genus Audibertia, the lower half of this connective 


3S 50D 


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 the middle, and 
surmounted by a one-celled anther, as shown in Fig. 511°. In 
Rosemary, the continuity is complete, although a minute reflexed 
tooth sometimes indicates the junction. 

470. Pollen, 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, 


512 515 54 51S 
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 are oblong; in the 


FIG. 508-511. Anthers, with upper part of filament. of several Labiate. 508. Of 
Monarda. 509. Ofa Calamintha. 510 Of two species of Salvia. with long and slender 
connective, the upper fork of which bears one anther-cell; the lower in @ (from Salvia 
Texana), bearing the othercell in an imperfect condition; in b (from S. coccinea), bear- 
ing none at all. 511. a. Same of Andibertia grandiflora, the lower fork of the connec- 
tive reduced to a naked spur; b, 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. 

FIG. 512-515. Forms of pollen: 512, from Mimulus moschatus; 513, Sicyos; 514. 
Echinocystis; 515, Hibiscus. 


POLLEN. eg 


Cichory and Thistle tribes, many-sided; in the Musk-plant, 
spirally grooved; in the Mallow family and the Squash and 


Pumpkin, beset with briskly projections, &c. The pollen of 
Pine, as well as that of the Onagracez, is not so simple, but 
appears to consist of three or four blended cells ; 
that of most Ericaceze 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 first 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 
Poti, sing. PoLirytum) of Orchises (Fig. 463), 
and the denser waxy ones of many 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, Circea; 
520. Kalmia; 521, Evening Primrose. 
FIG, 522. <A pair of pollinia of Asclepias. annexed by their caudicles to the gland. 


17 


258 THE FLOWER. 


nifying power of about three hundred diameters, is found to 
contain a multitude of minute particles (foville), 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 
water by 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 state. 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 coating tough. In 
many kinds of pollen, the grains, when immersed in water, soon 
distend to bursting, discharging the contents. 

474. Pollen-tubes. In others, and in most fresh 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 cleistogamous flowers (434), the pollen, while still in 
the anther, sends out its tubes, which may grow to a great length, 
in the mere moisture of the flower-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 Conifer, the grains of pollen have a peculiar internal structure or 
rather a development (suggestive of a homology with the microspores of some 
of the higher Cryptogamia), the contents at or before maturity undergoing 
division into two or three internal cells, only one of which acts in fertiliza- 
tion. When they act upon the ovule or are placed in water, and the inner 
coat swells by absorption, the bursting outer coat is commonly thrown off. 
In Pines and Firs (but not in Larch and Hemlock Spruce), the grain of 
pollen is singularly compound, consisting (as in Fig. 518) of a central arcuate 
body (the proper pollen-cell) bearing at each end an empty roundish cell. 
These are vesicular protrusions or appendages of the proper pollen-grain, of 
no known functional importance, except that they render such wind-dis- 
persed pollen more buoyant for transportation. 

2 Van Tieghem, in Ann. Sci. Nat. ser. 5, xii. 812, &e., 1869. 


ay 


GYNCECiIUM IN ANGIOSPERMS. 259 


unaltered until it is placed upon the stigma. The more or less 
viscid moisture of this incites a sim- = oe 2 
ilar growth, and also doubtless nour- A 
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 oyule, and upon a 
special formation within it. Consequent 526 

upon this an embryo is formed ; and the ovule now becomes a seed. 


Section VII. Tue Pistits, or Gynacium. 
§ 1. Ix 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 gyneecium 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 oyules 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.’ And plants with such gynecium 
are denominated ANGIOSPERMS or ANGIOSPERMOUS PLANTS. ‘To 
such only the present subsection specifically relates. 

477. The several terms which apply to the Gyneetum 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. 


260 THE FLOWER. 


enumerated and defined already (302, note): the elementary 
term is that of 

478. Carpel, Lat. CarpeLttuM. This is the term coined by Dunal, 
and is in common use. The better-formed word Carprprum 
(English Carpid) has been proposed, and best of all Carpo- 
PHYLLUM, in English Carpopiyll. For carpels are, as the word 
carpophylla denotes, pistil-leaves, or leaves of the gyncecium, 
7. e., seed-bearing or fructiferous phylla. They occupy the cen- 
tral or uppermost region of the flower. A carpel may be 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 homologue of a leaf. 

479. The morphological conception of an uncombined carpel 
is that of the blade of a leaf 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 style: 
some portion of this, usually the apex, not rarely a single or 
double line down the side which answers to the suture of the 
leaf-margins, and may be regarded as its continuation, is the 
stigma. ‘The carpellary leaf is always /zcurved: 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 be 
regarded as a portion of leaf- 
margins presented externally, des- 
Y titute of epidermis and formed 
be pot = of loose cellular tissue, which in 
anthesis is moist by some secretion. The ovwes are peculiar 
structures normally arising as outgrowths from the margins of 
the leaf, or some part of them, sometimes from the whole or 
a special portion of the upper or inner surface of the leaf. 

480. The carpellary leaf being involute, the suture, on which 
the ovules are normally borne, always looks toward the axis or 
centre of the flower. It is the only proper suture (or seam) a 
carpel can have. From its position it takes the name of /ener 
or Ventral Suture. And the opposite line or ridge, answering 
to the midrib of the leaf, being sometimes prominent and of the 


FIG. 527. A leaf incurving, to illustrate the morphology of a simple pistil or carpel. 
528. A carpel (of Isopyrum biternatum), cut across, the lateral stigma (here manifestly 
a double line) and the suture bearing the ovules turned toward the eye. 529. A ripe 
carpel of Marsh Marigold which has opened and shed the seeds: the points of attach- 
ment of the latter conspicuous along the edges of the carpel. 


GYNCCIUM IN ANGIOSPERMS. 261 
appearance of a suture, has been somewhat incongruously named 
the Outer or Dorsal Suture. 

481. The number of carpels in a gyneecium is simply expressed 
by adjective terms consisting of Greek numerals prefixed to this 
word: e. g., Monocarpellary, of a solitary carpel ; Dicarpellary, of 
two carpels; Zricarpellary, of three; Tetracarpellary, of four; 
Pentacarpellary, of five, and so on up to Polycarpellary, of many 
or at least of several and an indefinite number. Less general 
and only partially synonymous terms are such as Monogynons 
(of one pistil), Digynous (of two), Polygynous (of many), &e. 
These are adjective forms of the names of the orders, from 
Monogynia to Polygynia, in the Linnzean artificial classification, 
which either supposes the carpels to be separate or partly so, or 
confounds simple and compound pistils. 

482. When the gyneecium is of a solitary carpel, the position 
of this as regards the axis of inflorescence is not uniform; but 
commonly its back or dorsal suture is before the subtending 
bract, or in other words the ventral or oyule-bearing suture 
faces the axis of inflorescence. When there are two carpels, 
they face each other, bringing their ventral sutures into opposi- 
tion, and as to axis of inflorescence either median or transverse 
(291), but usually median, that is antero-posterior or in the 
line of bract and axis. Cruciferse, Capparidacez, and Fumari- 
acee are somewhat remarkable for having their two carpels 
right and left, that is, collateral or, in other words, transverse. 
When three, four, or a greater number, they divide the circle 
equally, or when numerous they take a spiral instead of verticil- 
late order, and occupy several or many ranks, as in Ranunculus, 
Magnolia, Potentilla, &c. 

483. The Gynecium may be either of separate carpels 
(Apocarpous), or of carpels coalescent into one body (Syncar- 
pous), or of all grades between the two. Apocarpous pistils are 
simple ; a syncarpous pistil is compound. 

484. In both, the essential parts are the ovary and the stigma. 
The style may be conspicuous and widely separate these two, 
as in Fig. 536-538 ; or hardly any, as in Fig. 532-535 ; or none 
at all, as in Fig. 530, 531, 533. 

485. Placenta. This name? is applied to any surface in the 
interior of the ovary on which ovules are borne. It has been 
stated (579) that these are usually borne upon the margins of 


1 Taken from aremote analogy with the placenta of the higher animals. 
The name appears to have been introduced into botany by Adanson. It has 
been termed Trophospermum or Spermophorum by some of the early modern 
botanists. 


262 THE FLOWER. 


ihe carpellary leaf, or upon some portion of what answers to 
them. When the ovules are numerous, and some- 
times when they are few, the combined leaf-edges 
enlarge to form a kind of receptacle for their attach- 
ment or support: this is the Placenta. In Fig. 530, 
the placenta is well developed, and also in such 
syncarpous ovaries as are illustrated in Fig. 536, 
537, 544, and 545. In very many others (such as 
Fig. 528, 531, 533), there is no particular enlarge- 
ment of the leaf-margins visible, and no particular 
ground for the use of this special term. Still it 
is commonly used, as occasion serves, even for the 
mere line or spot on which ovules are borne, as 
aie as for a more prominent development to which 
530 the name was originally applied. 

486. Simple or Apocarpous Pistils may be solitary, several, or 
numerous. When indefinitely numerous, they are seldom in one 
circle, but are capitate or spicate upon a proportionately enlarged 
or prolonged receptacle, as in Anemone, Ranunculus, and most 
strikingly in Myosurus; when reduced to a single one, as in 
Acta, Podophyllum,* Barberry, and Plum or Cherry, the car- 
pel mostly appears as if it were an actual termination of the 
floral axis. But even then the pistil 
is hardly ever quite symmetrical in 
shape: the ovary is somewhat gib- 
bous or unequal-sided (as in Fig, 
312,315, 316, 528, 531-533), and the 
stigma more or less oblique or even 
wholly lateral. The continuation of 

331 532 533 the latter down the whole length of 
the ventral side of the style (as in Fig. 528, and also Fig. 549) 
is not uncommon. In Schizandra (Fig. 531) it is continued 
downward on the ventral edge of the ovary as far as to its 
middle.* 


1 Abnormal specimens of Podophyllum peltatum are occasionally found 
having a gynecium of from two to six separate carpels. 

2 Pleurogyne, a Gentianaceous genus so named on this account, has no 
style nor apical stigma whatever, but has along stigma extending down the 
outside of each ovuliferous suture of its dicarpellary ovary for most of its 
length. 

FIG. 530. Single simple pistil of Podophyllum, cut across to show the placenta, &c. 

FIG. 531. Vertical section of a pistil of Schizandra coccinea; a side view showing 
the stigma decurrent down to the middle of the ovary. 532. Pistilof Hydrastis; ventral 


view. 533. Pistil of Actsea rubra, cut across, so as to show the interior of the ovary; 
ventral view. 


GYNGECIUM IN ANGIOSPERMS. 263 


487. As the placenta of a simple pistil belongs to the two 
united margins of the carpellary leaf, there is naturaily a double 
row of ovules, one to each margin. If the leaf- 
margins which are turned inward in the ovary be- 
low to bear the ovules are turned outward above to 
receive the polien (see Fig. 531), then the typical 
stigma should also be double or bilamellar. So it 
is seen to be in such carpels as those of Fig. 528, 
531-533, and indeed in very many stigmas of this 
class. Such division, or even a greater bifurcation 
of a monocarpellary stigma into two lobes or half- 
stigmas, is not anomalous. 

488. The ovary of a simple pistil should be 
untlocular, that is, should have a single cavity 
or cell (loculus), although, as will soon be seen, 
the converse does not hold true. Yet this cell in 
certain instances becomes bilocellate, being divided 
by a growth or intrusion from the back into two 
locelli. ‘This occurs more or less in the larger 
number of species of the Leguminous genus 
Astragalus, and the mode is shown in Fig. 534. 

489. Compound or Syncarpous Pistil.’| This consists of two. 
three, or a greater number of carpels coalescent into one body. 
A true compound pistil represents a whorl (in the simplest case 
a pair) of carpels united into one body, at least as to the ovary. 

490. The coalescence of a capitate or spicate mass of carpels 
or simple pistils of the same flower, imbricately heaped on the 
torus, as in Magnolia (Fig. 648) and Liriodendron, cannot 
properly be said to form a compound pistil. This heap of 
pistils may be called a Sorema. 

491. Morphologically, a compound pistil, as to the ovary, may 
be a pair or a circle of closed carpels or simple pistils brought 
into contact, and the contiguous parts united: this is illustrated, 
in Fig. 535-538. Or it may be formed of a whorl of open car- 
pellary leaves, joined each to each by the contiguous margins, 


5o4 


1 The terms apocarpous and syncarpous for pistils, the first of separate, the 
second of combined carpels, were introduced by Lindley. They have little 
advantage over the terms simple and compound. Moreover, the word 
syncarp or syncarpium had been appropriated to a sort of fruit of the class 
now called multiple, formed by the coalescence of several flowers, and also 
to that of a heap, head, or spike of carpels more or less cohering at matu- 
rity, as in a blackberry, or confluent in the flower, as in Magnolia. 

FIG. 534. Ovary or forming legume of Astragalus Canadensis, transversely divided, 


to show the false partition which, intruded from the back, divides the simple cell into 
two half-cells or locedii, 


264 THE FLOWER. 


in the manner of Fig. 542-545. Between these two there is 
every gradation. ‘The first forms a compound ovary, 

492. With two or more Cells and Axile Placente. For it is 
evident that, if the contiguous parts of a whorl of two or more 
closed carpels cohere, the resulting compound ovary should have 
as many cells as there are carpels in its composition, and that 
the placentze (one in the inner angle of each carpel) will all be 


585 


brought together in the axis of the compound pistil. And the 
partitions, termed Dissrpmrents, which divide the compound 
ovary into cells, manifestly consist of the united contiguous por- 
tions of the walls of the carpels. These necessarily are composed 
of two layers, one belonging to each carpel; and in fruit they 
often split into the two layers. True dissepiments and the true 
cells must accordingly be equal in number to the carpels of 
which the compound pistil is composed. That is, the ovary, or 
the resulting fruit, is bélocular or 2-celled, trilocular or 3-celled, 
quadrilocular or 4-celled, and so on, according to the number of 
dissepiments or cells. 

493. There may also be false dissepiments, mostly of the same 
character as that which in Fig. 534 divides the cell of a singie 
carpel. Such are found in Flax (Fig. 539-641), in Amelanchier 
or Service-berry, in Huckleberry (Gaylussacia), and in most of 


FIG. 535. Pistil of a Saxifrage composed of two carpels or simple pistils united 
below, but distinct above; cut across both above and below. 

FIG. 536. Pistil of common St. Johnswort, of three united ovaries; their styles 
distinct. 

FIG. 537. The same of another species of St. Johnswort (Hypericum prolificum), 
the styles also united into one, which, however, may split apart in the fruit. 

FIG. 538. Pistil of Tradescantia or Spiderwort, even the three stigmas united into 
one. The ovary in all cut across to show the internal structure. 


vet ie 


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. 


on) 540 Hl 


494. On the other hand, even the true dissepiments which 
belong to such a compound ovary may be abortive or evanescent, 
the placentze remaining in the axis combined into a column. 
(499.) The second modification of the compound pistil (491) 
normally has an ovary, 

495. With one Cell and Parietal Placente. That is, the 
placentz 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 placenta. 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 placente. 

FIG. 544. Transverse section of the ovary of Hypericvm graveolens; the three large 
placenta meeting in the centre, but not cohering. 545. Similar section of a ripe capsule 
of the same; the placentze now evidently parietal 


256 THE FLOWER. 


of a congenital development of organs in union which, in the 
development of a vegetative shoot, would be leaves.* This case 
is represented by the combination of open carpellary 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. 256, 
257. Every gradation is found between axile and parietal 
placentation. Sometimes the placentz are strictly on the pari- 
etes or wall (Fig. 545, 547); sometimes borne inwards on 
incomplete dissepiments (Fig. 548); and sometimes they are 
brought firmly together in the axis, as in Fig. 544, though sepa- 
rable, and indeed separated in the fruiting stage. _ 

496. A compound ovary with parietal placentz is necessarily 
one-celled (unilucular) ; except it be divided by an anomalous 
partition, such as is found in Cruciferz (Fig. 395) and in many 
Bignoniacee. 

497. Normal placentz are necessarily double: when parietal, 
the two halves belong to different leaves ; when axile, to the same 
leaf. These two halves may diverge or be widely separated, 


sometimes even at their origin, as in Aphyllon and some other 
Orobanchacez, in which a dicarpellary ovary has four almost 
equidistant placentze; or in such cases the placentz may be 
regarded as intra-marginal instead of marginal. 

498. The placentz of a two-several-celled ovary, such as in 
Fig. 536, 537, &e., may be described in the plural number, 
being one in each carpel: or when consolidated into a central 
column, and well covered with ovules, they may be said to form 
one (compound) placenta. Then when the dissepiments early 
disappear, or are abortive from the first, the result is a compound 
ovary of this class, 

499. With one Cell and Free Central Placenta. In Caryo- 
phyllacee (Fig. 549, 550) and Portulacacez, this evidently 
results from the obliteration of the dissepiments (as many as 
there are styles or stigmas), vestiges of which may be sometimes 


FIG. 546. Diagram (ground-plan) to illustrate free central placentation produced 
by abortion of dissepiments. 547. Same of strict parietal placentation. 548. Same with 
the placentz carried inward on imperfect dissepiments. 


GYNGCIUM IN ANGIOSPETMS. 267 


detected, while certain plants of the same familics, 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 \ 
p 


549 


bottom, and the placentze there conspicuously devel- 
oped and completely united. The basal placenta- 
tion of Dionzea is unavoidably so explained, its 
nearest relative, Drosera (Fig. 553), having parietal 
placentze. And this leads to a probable explanation 
of the case in Primulacez, where a large free central 
placenta fills the centre of the cell, and no trace of 
dissepiments can be detected.’ 

501. The idca maintained in former editions is 
still adhered to; namely, that placentz belong to 
cearpels 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.? 

502. Ovules cover the whole internal face of the carpels in 
Butomus and its relatives, also of the Water-Lilies (both Nym- 
pheea and Nuphar, Fig. 551) excepting the inner angle, to which 
they 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. 529 (1869) ; 
Celakowsky, Vergleichende Darstellung der Placenten, &c. (1876) ; Wariing, 
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 Plante Javanice Rariores, 107-112. Schleiden, End- 
licher, and others took the opposite view, 7. 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 he gives his entire 
adhesion to this conclusion) ; also Warming’s memoir, De l’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, 


268 THE FLOWER. 


margins of the carpellary leaf. In many species of Gentian, as 
also in Obolaria and Bartonia, of the same 
family, the whole internal face of a dicar- 
pellary ovary is thickly ovuliferous. 
503. Perhaps the parietal placentae in 
Parnassia (Fig. 552) are borne on 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 Papaveracez, also in some Cruciferee ; 
and in some of the cases each stigma 
is more or less two-lobed. ‘This sug- 
gests the explanation,’ here probably 
the true one, which supposes that the 
placentze are borne on the leaf-margins 
in the normal way, but that each 
stigma is two-parted (as if the carpel- 
lary leaf were deeply notched at the 
apex, and so its two stigmatic leaf- 
margins separate, as Drosera illus- Ss 
trates, Fig. 553), and that the two oy Soa 
half-stigmas of adjacent carpels have 338 
coalesced into one body, which would 
of course stand over the parietal placentze beneath. Each stigma 
in such a case, as well as each parietal placenta, would consist 
of the united margins of two adjacent carpels. 


§ 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 flowers, 
these plants are of a low or simplified type, in some respects not 
obviously homologous with the Angiosperms which now consti- 
tute the immense majority of phanogamous plants. But, up to 
a comparatively late geological period, Gymnosperms appear to 
have been the only flower-bearing plants. They are represented 


1 Given by Brown, in the Plante Javanice Rariores, above referred to. 


FIG. 551. Transverse section of an ovary of Nymplza odorata, the carpels ovulifer- 
ous over the whole interior surface. 

FIG. 552. Pistil of Parnassix, with ovary transversely divided. 

FIG. 553. Pistil of Drosera filiformis, 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 they cannot be illustrated by a common description. 
The largest order, Conifers, is familiar, and contains a good 
share of the most important forest trees of temperate climates. 
The smallest, Gnetacez, chiefly tropical or of warm regions, 
lies between Gymnosperms and common Dicotyledons. The 
third, Cyeadaceze, 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.’ 

505. In Gnetaceer, Gymnosperms and Angiosperms almost 
come together. The flowers have a perianth (diphyllous or 
tetraphyllous) ; the stamens have a distinct filament and anther ; 
and the gyneecium is a sac (presumably of two carpophylls) 
open at the top and filled at bottom by a single oyule of the 
simplest kind, 7. 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 thealmost hermaphrodite sterile flower of Welwitschia, 
this takes the form of a much dilated stigma, which is even beset 
with seeming stigmatic papille. 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.* 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 morphoiogy still in part 
unsettled, need not here be given. The history and the idea of gymnospermy 
began with Robert Brown’s paper on Kingia, “ with Observations .. . . on 
the Female Flower of Cycadeze and Conifer,’ 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 Conifere. 

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 FLOWER. 


genus Gnetum, shrubs or trees, with nearly the aspect of 
Angiosperms, having broad and pinnately-veined leaves; Wel- 
witschia of tropical W. Africa, remarkable for its persistent 
cotyledons which form the only foliage of a woody and long- 
enduring plant, and for its stem or trunk which broadens with- 
out lengthening, except in its flower-stalks ; also Ephedra, of 
much branched shrubs, mainly of warm-temperate regions, leafiess 
or nearly so, one species of which inhabits Europe and two the 
southern borders of the United States. 

506. The flowers in all Gymnosperms are diclinous, either 
dicecious or moneecious ; except that those of the strange Gneta- 
ceous genus Welwitschia are structurally polygamous, the male 
flowers having a well-formed but sterile gyncecium. 

507. In Coniferx, the largest and most important type, are 
embraced all the familiar Gymnosperms of temperate regions, 
Pines, Firs, Cedars, Cypresses, which > 
bear their flowers in catkin-like clusters {Y= 
and their fruit in cones, and also the \¥,. 
Yews and allied trees which do not/C-~ 
produce cones. Perianth being want- 
ing and the sexes wholly separate, the floral type 
is so degraded that it becomes doubtful whether 
each cluster of anthers, or of ovuliferous seales 
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 flower. It is here 
assumed that each stamen ( 


of the one and each oyu- 
liferous scale of the other 
answers to a flower of the 
simplest sort.! The anthers 
oS are extrorse, the cells or 
pollen-sacs belonging to the outer or lower side of a scale or a 


557 


1 Tt will be seen that, for the female flowers, this follows of course from 
generally accepted view ; and, where this is conceded, analogy may extend it 
to the male catkins also: yet in such cases, where all the phylla of an indefiniie 
simple axis are stamens, spirally arranged on it, the difference betwecn 
inflorescence and male flower completely vanishes. 


FIG. 554. Female flower of a Yew, an ovule surrounded by its bracts. 555. Longi- 
tudinal and more enlarged section of a female flower of Yew and of the upper part of 
the shoot it terminates: the thick coat of the ovule open at the top, the nucleus within, 
and the beginning of the disk outside of the coat, are seen in section. After Stras- 
burger. 

FIG, 556. Young fruit (berry-like cup surrounding the seed) of Yew. 557. “Longi- 
tudinal section of a mature fruit of the same. After Decaisne. 


GYNGC!IUM OF GYMNOSPERMS. 271 


connective: sometimes these sacs or cells are two, and the organ 
evidently homologous with an ordinary stamen: often they are 
more numerous (from e 
three to twenty) and 
yariously disposed. 
508. The Yew Fam- 
ily (Taxinez) is next to 
Gnetacez in structure. 
It is generally ranked as : 
a suborder of Coniferze, 559 
but it may claim to be a distinct order. The gynecium is a 
naked ovule, terminating a stem,’ 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 
gynecium of Torreya, except that the cup- 
shaped disk develops almost simuitaneously 
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. 
598), 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 ovyule 


1 Tt does not therefore follow that the ovule isa 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 adian ifolia. 5584. 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 ernustaceous seed- 
coat, within which is the keruel of the seed; at the base on one side a sterile ovule is 
seen. After Decaisne. 

FIG. 560. Female flower of Podocarpus (an oyule 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. 


212 THE FLOWER. 


is inverted on a more or less lengthened and stout support, which 
is conceived to represent the carpel. (Fig. 560, 561.) 

509. In the true Coniferze, to which Pines, Cypresses, and all 
such cone-fruiting trees belong, the ovules are borne on or in 
the axils of scales which are imbricated on a simple axis, in a 
spicate or capitate manner; and the male flowers, each a single 
stamen, are also similarly spicate or capitate. Both are com- 
monly termed aments or catkins; and the female 
ones properly so, according to the present view ; 
but the only scales of the male catkins are parts 
of the anther, being a dilated tip of the counective 
in Pines, and a scale bearing anther-cells or pollen- 
sacs on its back in Cypress. 

510. In the Pine tribe the flowering female catkin consists of 
bracts, spirally imbricated on the cauline axis: in the axil of 
each bract or sterile scale is developed a scale 
which bears two ovules, and is therefore regarded 
as of carpellary nature. These ovules are pro- 
duced on the lower part of the upper face of this 
carpellary scale, and are wholly adherent to it 
quite to the orifice, which is directed downward. 
(Fig. 562, 563.) The ovuliferous scale in 
becoming fructiferous usually much and soon out- 
grows the bract, which is concealed in the Pine- 
cone (or sometimes obliterated) ; but it remains 
conspicuous in sundry Fir-cones. After fertil- 
ization, the scales, successively covering each 
other in close imbrication, protect the growing 
seeds as effectually as would a closed ovary. 
Sooner or later after ripening the scales diverge, and the seeds 
peel off the face of the scale with a wing attached, and fall or 
are dispersed by the wind.” 


1 Among those who admit as well as those who reject gymnospermy, 
there has been much controversy over the morphology of the parts. With 
the former, the discussion turns on the character of the ovuliferous scale. 
As to this, the hypothesis originally proposed by Mohl, and adopted by 
Braun, is now said to be satisfactorily demonstrated by Stenzel, in Nov. Act. 
Nat. Cur. xxxviii. 1876. See note by Engelmann in Amer. Jour. Sci. Dec. 
1876, and also the preface to the second part of Eichler’s Bliithendiagramme, 


FIG. 562. View of the upper face of a carpellary scale of a Larch, showing the pair 
of adnate ovules. 

FIG. 563. Similar view of a carpellary scale of a Larch, and of a bract behind if. 
564. Ground plan of the same in diagram, reversed; the upper figure denoting the axis 
of the cone, the lower the bract, the middle one the carpellary scale and the two ovules 
borne on its face. After Eichler. 


GYNCECIUM 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 Cypress or 
so-called Cypress of the Southern United States. 

512. In the true Cypress tribe (Cupressinez) the cone-scales, 
which are never numerous, are opposite or verticillate, 7. e. like 
the foliage-leaves, in whorls of twos, threes, or 
sometimes fours; and the ovules are from two to 


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 dors:/ 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 phenogamous plants, but is 
the rule in Ferns, from something like which Coniferee 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 Araucaria imbricata. After Eichler. 

FIG. 566. Branchlet of the American Arbor-Vitz, considerably larger than in na- 
ture, with a forming fertile cone. 567. One of the scales remoyed 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 


274 THE FLOWER. 


several at or on the base of each cone-scale, always with orifice 
upward. Arbor-Vite (Fig. 566, 567) has a single paix of ovules 
to the scale; Junipers, sometimes only one; true Cypresses (as 
in Fig. 568), often a dozen or more. At flowering time, the cone- 
scales mostly appear 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.? 


513. In Cyeadacee, the type of the flower of Angiosperms is 
almost or quite lost; yet the organs may be homologized with 
those of Coniferze, which these plants are wholly unlike in habit. 


1 This internal and ovuliferous scale may seem to be wholly hypetheti- 
cal, and assumed to homologize the cupressineous with the abietineous 
cone. Without it, we should have to consider that, while in Abietinee the 
ovules belong to leaves of a secondary axis, in Cupressineew they are borne 
on those of a primary axis, or else are axillary productions without carpels. 
But in the Araucaria tribe the internal scale is obvious ; and there are sufii- 


FIG. 569-575. Zamia, chiefly Z. media, after Richard. 569. A male plant. 570 
Lower part of a male catkin. 571. A stamen removed, showing numerous small pollen- 
sacs under the peltate top. 572. A female catkin, with a quarter section cut away. 
573. A female flower or carpel, with two enlarging ovules or young seeds. 574. Ripe 
seed, with the thick fleshy coat cut away at apex. 575. Longitudinal section of ripe 
seed, more enlarged. 


GYNGCIUM OF GYMNOSPERMS. 273 


Their likeness to Palnis 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 they any 
further resemblance to Ferns, except that in some the leaflets are 
circinate 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 


576 577 578 


the United States, and a more striking one (Cycas revoluta, well 
known in cultivation) in the southern parts of Japan. 

514. Following the analogy of Conifer, each scale (whether 
of the pollen-bearing or the oyule- 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 to 
induce the adoption of it by Parlatore, who rejected the idea of gymnospermy ; 
and, finally, this composition is nearly demonstrated by Van Tieghem (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. 


nally much alike in the two sexes, which throughout the family 
occupy separate plants. The male flower (Fig. 568) or stamen, 
if it may be so termed, bears indefinite pollen-sacs on the under 
side of the peltate portion, sometimes extending to che upper 
part of its stalk. The homologous female flower, or carpophyll, 
bears a suspended ovule on each side of the stalk (Fig. 573), 
which becomes a large flesny-coated seed. In Cycas the male 
ament is not very dissimilar, although on a larger scale. But 
the carpophylls are evident leaves, not condensed into an ament, 
but loose or spreading, of a character and aspect intermediate 
between the lax bud-scales which precede and the pinnate foliage- 
leaves which follow them in development. Along the margin 
of what would be leaf-blade they bear ovules in place of leaflets, 
lobes, or teeth (Fig. 576-578) ; and these, when fertilized from 
the male flowers, mature into large and drupaceous naked seeds. 
Even without fertilization, such seeds grow to their full size on 
the female plant of the common Cycas (or falsely so-called Sago 
Palm), but form no embryo. 


Section VIII. Tue Ovoute.! 


515. Ovules (302) are peculiar outgrowths or productions of 
earpels which, upon the formation of an embryo within, become 
seeds. In the angiospermous gynecium (476) they are nor- 
mally produced along the margins, or some part of the margins, 
of the carpellary leaf (478), either immediately, or by the in- 
termediation of a placenta (485), 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 the 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 Gymnosperms (504-514) the ovules 
are borne on the face of the carpellary 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.) 

516. As to attachment. ovules are either sessile, 7. e. stalk- 
less, or on a stalk of their own (Fig. 582, 584), the FuNicuLUs 
or Poposperm. As to number they are either solitary, few, or 
en ee Se ee 


1 Lat. Ovulum, pl. Ocula, diminutive of ovum (egg), perhaps first used by 
Adanson 


OVULES. ZT 


indefinitely 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 ereef, 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, 
as in Fig. 581. 

518. The body and only essential part of an ovule is its 
Nuctevus. This in most cases is invested by one or two proper 
coats. The coats are sacs with a narrow orifice, the ForaAMEN. 
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 Exosrome, of the inner ENDOSTOME ; 
literally the outer and the inner 
orifice. The coats themselves have 
been named Primrye and SrEcun- 
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 Cuataza. The attachment of the 
ovule to its funiculus or support, which in the seed becomes the 


379 


J Hs 


582 oS4 


FIG. 579. Ovary of a Buttercup, divided lengthwise, to display its ascending 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. 5824), showing the outer coat, a, the inner, b, the nucleus, c, the chalaza, or place 
of junction of these parts, d. (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. An amphitropous oyule. 


* 278 THE FLOWER. 


HixvM, takes also this latter name in the ovule. In the simplest 
form of ovule (as in Fig. 582, 580), hilum and chalaza are one. 
So also in cases where the body of the ovule incurves, as in Fig. 
583. But very commonly the place of attachment, which becomes 
the hilum, is more or less distant from the chalaza ; as in Fig. 584 
and 587, where the hilum is lateral, but the chalaza at the larger 
end, the two being connected by a short ridge; and in Pig. 588 
the two are separated by the whole length of the ovule. 

519. The simplest and most rudimentary oyule is that with- 
out a coat, as in Mistletoe and the whole order Loranthaceze, and 
in Santalacezee and Gnetacee. 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, 7. e. to 
uncovered ovule, not to uncovered nucleus. The oyule consist- 
ing only of nucleus may be termed (after Alph. DeCandolle) 
simple, or better achlamydeous.* 

520. The tunicated or chlamydeous oyule 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.” 

521. Orthotropous (Fig. 580, 582, 585), or straight ovule, is 
the simplest but least common species, being that in which the 


chalaza is at the evident base, and the orifice at the opposite 
extremity, the whole ovule straight (as the first part of the name 
denotes) and symmetrical. Atropous, meaning not turned at 
all, is a later and etymologically much better name, but it has 


1 An epidermal stratum or tegument may not be wanting to such ovules, 
forming a sort of adherent covering; but this in nature and origin is not 
similar to the ovular coats. 

2 In Latin form, orthotropa, campylotropa, anatropa, amphitropa, — names given’ 
by Mirbel, and referring to the way in which the ovule is turned either on 
itself or on its support. Some English botanists incongruously write ortho. 
tropal, campylotropal, &c. 


FIG. 585. Orthotropous or Atropous ovule of Buckwheat. 586. Campylotropous 
ovule of Chickweed. 587. Amphitropous ovule of Mallow. 588. Anatropous ovule of a 
Violet. The letter k indicates the hilum; c, the chalaza, which in 585 and 586 corre- 
sponds to the hilum; /, the foramen or orifice; 7, the rhaphe. 


OVULES. 279 


not come into general use. This ovule is characteristic of 
Polygonacez, the proper Urticaceae, Cistacer, &e. 

522. Campylotropous (Fig. 585, 586) 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 
campylotropous develops unequally, one side enlarging much 
more than the other, especially at the base, until the ovule 
becomes reniform, and chalaza and orifice are brought into 
close proximity. Campylotropous ovules are characteristic of 
Cruciferze, Capparidaceze, Resedaceze, Caryophyllacez, and 
Chenopodiacez. 

523. Amphitropous (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 hilum 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 Primulacez, and 
is common in Legumminos. 

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, 7), is named the Ruapure. This is not 


280 THE FLOWER. 


at all a seam, 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 oyule is the nucleus (518), 
or the nucleus surmounting a rudimentary funiculus, 
The nacleus is soft cellular tissue only, from first to 
last. The achlamydeous ovule (519) undergoes no 
further development except in size or shape. Indeed 
sometimes (as in Balanophoreze) 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 fibro-vascular tissue, 
especially spiral ducts, may be found in the funiculus and cha- 
laza, sometimes extending into the coats. 

528. 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. 


FIG. 589. Magnified view of a vertical section of a carpel of Magnolia Umbrella, 
about a month before anthesis, showing one of the two nascent ovules, at this time 
only nucleus. 

FIG. 590-597. Further development of the ovule of Magnolia Umbrella, showing the 
formation of the coats and the anatropy. 590. Ovule a week older than in 589. 591. 
Same a week or two later. 592. Same a few days later 593 Same from a nearly full- 
grown flower-bud. 594. Same at time of anthesis. 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 oyule 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 
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 (7. 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 By comparison of Fig. 578 with 576 and the like, it may be perceived 
that the difference is explicable by 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 cf 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 Pl. Javan. Rar. 151), and confirmed by the analyses of 
Sprague (Gray, Gen. III. ii. 168, plates 163-169), 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. Pl. 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 (b), nucleus (ce), 


and the bundle of spiral ducts (() in the rhaphe (running from placenta to chalaza) 
indicated. 


282 THE FLOWER. 


530. Origin and Nature of the Ovule. It has been already 
stated in general terms that ovules are peculiar outgrowths 
or productions, generally of the margins of carpellary leaves 
(515) ; that they are composed of parenchymatous cellular sub- 
stance, at least as to the nucleus, of which the simplest ovule 
wholly consists (526) ; that the coats originate subsequently to 
the nucleus ; and that the outer coat is of later origin than the 
inner one. (518.) The mamiliform protuberance of which the 
forming ovule at first consists originates in one or more cells of 
a layer directly beneath the epidermis.* 

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 leayes, 
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 


apotropous, epitropous, and heterotropous (the first two new, the last employed 
in a new sense), the more so since the application is confused with hypo- 
thetical considerations and the necessity of bringing the ovules ideally back 
to ascending or horizontal positions. It may be stated, briefly, that /Zetero- 
tropous, in 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 with its 
rhaphe next the placental axis, and a hanging one has its rhaphe averse 
from it; /pitropous, when an erect or ascending ovule has its rhaphe averse, 
and a hanging one has it adverse. 

1 Hofmeister’s statement that the simple ovule of Orchis originates in 
the division of a single epidermal cell (and is therefore a trichome) is con- 
troverted by Strasburger 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, De l’Ovule; 
also the papers of Celakowsky, Van Tieghem, &c., referred to in notes to 
paragraphs 500, 501. 


OVULES. 283 


ereat advantage of this view is that it serves to homologize the 
fructification of Flowering Plants with that of the higher klower- 
less Plants, or the Ferns, the sporangia or analogues of the 
ovule being outgrowths of the leaf.! 

532. Origination of the Embryo. ‘The whole process of fer- 
tilization and the resulting produc- ee 
tion of the embryo, also the history Y a 
of the subject, belongs to the suc- 
ceeding volume, involving as they 
do questions of minute anatomy and 
of physiology. But a general idea 
may here be given of the way in 
which the embryo originates. The 
tube which a grain of pollen sends 
forth into the stigma (074, 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 embryo-sac forms a globule, and this at the time 


! The advocates of this view naturally maintain that ovules and placente 
always belong to leaves, and never truly to a cauline axis; that in the pre- 
central placentation of Primulacez, 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 carpophyll, 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 stigmas, 
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 (0), and reached the 
embryo-sac (s) near the embryonal vesicle (7). 


284 THE FLOWER. 


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 
599 600 601 wall of the sac exactly 
opposite the termination 
of the pollen-tube. This 
is called the embryonal 
vesicle. To it the con- 
tents of the pollen-tube 
are in some manner trans- 
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 swspensor or pro- 
embryo). The terminal cell of this divides again and again in 
three directions, producing a mass of cells which shapes itself 
eos 605 606 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 
micropyle 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, Santalum, &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 Strasburger, Ueber Polyembryonie, in Zeitschr. Naturwis. Jena, xii. 
1878 (see Amer. Jour. Sci. April, 1879). 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 at its extremity. 600. 
The same, with the embryo a little more developed. 601. The same, more developed 
still, the cotyledons faintly indicated at the lower end. 602. Same, with the incipient 
cotyledons more manifest. 603. The embryo nearly completed. 

FIG. 604-606. Formingembryo froma half-grown seed of Buckwheat, in three stages. 
607. Same, with the cotyledons fully developed. 


THE FRUIT. 285 


anomalous reproduction are therefore now known, which 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.’ The production of bulblets in place of 
seed or embryo answers to this in Flowering plants. 

Parthenogeny, the counterpart analogue of apogamy, 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. 


GHAPTER VIL. 
THE FRUIT. 


Section I. Irs 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 ovary, to a cluster of such ovaries, at least 
when somewhat coherent, to a ripened ovary with calyx 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 
Celebogyne offers an exception. The female of this dicecious 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 embryo 
thus formed is adventive, the embryonal vesicle perishing. Purthenogenesis, 
of which Czlebogyne 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. ix.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 during fruc- 
tification, either by the abortion or obliteration of parts, or by 
accessory 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 
matured, 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, there 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 bilocellate, 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 five-celled ovary of common Flax early becomes spuriously 
ten-celled (morphologically speaking, not 10-locular, but 10- 
locellate), by a false partition extending from the back of each 


FIG. 608. Longitudinal section of the ovary of a Buckeye (sculus Pavia), showing 
the pairs of ovules in two of the cells. 609. Transverse section of the same displaying all 
three cells and six ovules. 610. Same of half-grown fruit, with single fertile seed, 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 
a maturing ovary (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. 625-627), 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 Borraginacez) , 
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 ovary, 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 Compositz 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 frait, 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 
ovary 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 
earpels of Columbine, and Marsh Marigold (Caltha), or the pod 
of Colutea or Bladder Senna. In others it thickens and becomes 
at maturity either dry throughout, as in nuts and capsules; or 
fleshy or pulpy throughout, as in berries; or hard-rinded with- 


288 THE FRUIT. 


out but soft within, as in a pepo; or fleshy or berry-like without, 
but indurated within, as in all stone-fruits, such as the cherry 
and peach. 

541. When the walls of a pericarp consist of two layers of dis- 
similar texture (as in a peach) the outer layer is called Exocarp, 
the inner Enpocarp, 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 Ericarp. When it is 
fleshy or pulpy it is named Sarcocarr. When the endocarp within 
a sarcocarp is hard and bony or crustaceous, forming a shell or 
stone, thisis termed a PurameN. When three concentric layers are 
distinguishable in a pericarp, the middle one is called Mresocarp. 

042. 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; cndehiscent 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 many seeds mostly dehiscent. Seeds pro- 
vided with a wing or coma or any analogous help to dispersion 
are always in indehiscent pericarps. Permanently fleshy peri- 
carps are indehiscent, stone-fruits as well as berries. But in 
some stone-fruits (7. e., with indurated endocarp and fleshy 
exocarp), such as those of Almond (Fig. 640) 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. Dehiscence, the opening of a pericarp for the discharge 
of the contained seeds, is ~-gular or irregular; 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 Pop. 

544. Regular 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 immer, ven- 
tral, or ovuliferous 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 septierda/, that is, 
as the term denotes, cutting through the septa or dissepiments ; 
and the loculicidal, that is, cutting into the /oculi 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 
away on the edges of the par- 
titions or introflexed valves ; 
also by Rhododendron, Kal- 
mia, and the like, in which 
the placentee 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 {| © Z 
not of itself open the cells. 
Such separated carpels when 

aa one-seeded not rarely remain “_|_~ 
closed, as in Mallow, Ver- op 
bena, &e. Or when dehiscent they may open both by the ventral 
and dorsal sutures; 7. 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 dehiscence. 

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, G£nothera, &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. Septifragal dehiscence, ¢.e., a breaking away of the 
valves from the septa or partitions, as shown in Fig. 616. This 
represents the loculicidal form of the septifragal mode, which is 
less common than that of the accompanying diagram, Fig. 617. 


Here the partitions alternate 

Pa YN __ With the valves; that is, the’ 

dehiscence of the pericarp is 

of the septicidal order, as 

near as may be, but the par- 

titions do not split, wherefore 

the valves break away at the 

ae mm common junction. To this 

the term marginicidal has been applied. It occurs in the 2-3- 

carpellary capsule of Ipomeea (especially in the common Morning 

Glory), in the 5-carpellary capsule of the North American species 

of Bergia; likewise in the 2-carpellary pod of Crucifere (Fig. 

623), with a difference that the placentz from which the valves 
break away are here parietal and the partition is abnormal. 

551. The terms septicidal and loculicidal apply equally in plan, 
though not with etymological correctness, to one-celled capsules 
with either parietal (495) or free central (599) placentee. When 
the dehiscence is of the septicidal type and the placentation pari- 
etal, the (half) placentze are borne on the margin of the valves, 
as in the Gentian family and the species of Hypericum with one- 
celled capsule. When the placentz are borne on the middle 
of the valves, as in Violets, the dehiscence is of the loculicidal 
type. In the case of free central placentze with no trace of 
partitions, the character of the dehiscence may usually be deter- 
mined by the position of the styles or stigmas relative to the 
valves. 

552. Dehiscence may be quite normal although very partial, 
as when confined to the apex of the capsule of Cerastium and 
of Primula, and even to the pores under the radiate stigmas 
of Poppy. 

553. Irregular or abnormal dehiscence is such as has no respect 
to the normal sutures; as where the dehiscence is transverse ; 


FIG. 616. Diagram of loculicidally septifragal dehiscence. 617. Same of septicidally 
or rather marginicidally septifragal dehiscence. 


a 


ITS KINDS. 291 


either extending part way round, as in the pod of Jefiersonia, 
or completely round, so that the upper part falls off like an 
unhinged lid. This ezrewmscissile dehiscence occurs in many 
plants of widely different orders ; such, for example, as Purslane 
(Fig. 621), genuine Amaranths, Plantain, Pimpernel, and Hen- 
bane. In other cases, as in Antirrhinum (Snap-dragon) and 
its allies, the cells burst by irregular laceration at a definite 
point, and discharge the seeds through the ragged perforation ; 
or one or more neat valvular orifices are formed on some parts 
of the wall, as in Campanula. 


Section II. Tue Krinps or Frutr. 


554. Fruirs have been minutely classified and named ;! but 
the terms in ordinary use are not very numerous. <A rigorously 
exact and particular classification, discriminating between the 
fruits derived from simple and from compound pistils, or between 
those with and without an adnate calyx, is too recondite and 
technical, and sometimes too hypothetical, for practical pur- 
poses. It is neither convenient nor philosophical to give a 
substantive name to every modification of the same organ. For 
all ordinary purposes, both of morphological and systematic 
botany, it will suffice to characterize the principal kinds under 
the four classes of — 

Simple fruits, those which result from the ripening of a single 
pistil ; 

Aggregate, those of a cluster of carpels of one flower crowded 
into a mass; 

Accessory or Anthocarpus, where the principal mass consists 
of the surroundings or support of either a simple or an aggregate 
fruit ; 

Multiple or Collective, formed by the union or compact aggre- 
gation of the pistils of several flowers, or of more than one, 

555. Simple Fruits may be distinguished, upon differences of 
texture, into Dry Fruits, Stone Fruits, and Baccate Fruits; or, 
better, into Dry and Fileshy ; and the first may be divided into 


1 The greater part of the forty-three substantive names of Desvaux’s, 
and even of the thirty-six of Dumortier’s and of Lindley’s elaborate classi- 
fications of fruits have never found employment in systematic botany, and 
doubtless never will be used. Yet a detailed carpological classification has 
its uses for the student. Among the more recent attempts are the successive 
ones of Dickson, McNab, and Masters. See Nature, iv. 347 (also in Trimen’s 
Jour. Bot. 1871, 310), iv. 475, and v. 6. 


292 THE FRUIE. 


dehiscent and indehiscent kinds.! Theoretically, each kind may 
be divided into those of a simple and those of a compound pistil, 
and some would make the primary division on this character. 
Some also would separate fruits with adnate or superior calyx 
from those free of all such combination. But in practice these 
differences can seldom be indicated by substantative names. 
The name of berry is equally applicable to the fruit resulting 
from the single carpel of Actza, the syncarpous ovary of the 
grape, and the similar ovary with adnate calyx of a gooseberry 
and cranberry. It should be, understood that the kinds shade 
off one into another most freely. 

556. Dehiscent Fruits (543), or Pods, are distinguishable into 
apocarpous, or of single carpels, and syncarpous, of more than 
one earpel, 7. e. the first of a simple, the second of a compound 
pistil. The first kind is mainly represented by the olliele and 
the Legume; the second, by the Capsule and its modifications. 

557. A Follicle is a pod formed 
of a simple pistil, and dehiscent by 
one suture (this almost always 
the ventral or inner suture) alone ; 
as in the Larkspur, Columbine, 
Peony, and Marsh-Marigold (Fig. 

sa 618) ; also in Milkweed and Dog- 
bane. There may be several follicles or only 
one to a flower, even in the same genus, as in 
Larkspurs, Cimicifuga, &c. In Magnolia 
(Fig. 648-650), fleshy carpels become follicles 
dehiscent by the dorsal suture. 

558. A Legume is the pod formed of a 
simple pistil which is dehiscent by both sut- 
ures (as in the Pea, Fig. 619), so dividing 
into two pieces or valves. (544.) This is the 
fruit of the Pulse Family, accordingly named 
Leguminosze (Leguminous plants): indeed, 
the name of legume is restricted to the fruits of this family, 
and in descriptive botany is extended to all the modifications 


1 Dr. Masters’s modification of Dickson’s and McNab’s classification of 
simple fruits, as to primary kinds, is into 

1. Nuts, or Achcenocarps, dry and indehiscent ; 

2. Pods, or Reqmacarps, dry, dehiscent ; 

3. Stone-fruits, or Pyrenocarps, fleshy without, indurated within, indehiscent ; 

4. Berries, or Sarcocarps, fleshy throughout, indehiscent. 


FIG. 618. A dehiscent follicle of Marsh-Marigold, Caltha palustris. 
FIG. 619. Legume cf a Sweet Pea, already dehiscent. 620. Loment of a Desmodium, 


ITS KINDS. 293 


which that order presents. Some of these, in fact, are in- 
dehiscent and reduced to akenes; some break up at maturity 
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 Lomenr, 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 Repium, 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 valves. ‘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; @. ef the dehiscence 
is circumscisstle. (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. on 

561. A Silique is a narrow two-valved capsule, with two pari- 
etal placentze, 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 placente, 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 Capsella or Shepherd’s- 
Purse, lateral view, and an oblique view of the same with one valve removed. 


294 THE FRUIT. 


ing the pod two-celled in an anomalous manner. A SILIcLe 
(Silieula, diminutive of siliqua) is merely a short silique, the 
length of which does not more than twice or thrice surpass the 
breadth ; such as that of Shepherd’s-Purse (Fig. 624). and of 
Lunaria, Candytuft, &ce. 

062. Indehiscent Dry Fruits are almost always one-seeded or 
very few-seeded. If numerous, the seeds thus placed would not 
be dispersed. The ordinary kinds are strictly one- 
seeded, and in common language are often con- 
founded with seeds. The ways in which such fruits 
are dispersed are various. In the following case, 
the adaptation of the pericarp to dispersion by wind 
distinguishes the species of fruit. 

965. The Samara, sometimes called in English a 
na is an indehiscent one-seeded fruit provided 
with a wing. In the White Ash 
the wing is terminal (Fig. 625) ; 
in other species the whole fruit 
is wing-margined ; in Birch and 
Elm (Fig. 626) the wing sur- 
rounds the body of the pericarp ; 
and the Maple fruit is a double 
samara or pair of such fruits, con- 

626 spicuously winged from the apex. 2 

564. Akene (Lat. Achenium) is a general name for all the 
une-seeded, dry and hard, indehiscent and seed-like small fruits, 
such as are popularly taken for naked 
seeds. But that they are true pistils, 
or ovaries ripened, is evident from the 
style or stigma they bear, or from the 
sear left by its fall; and a section 
brings to view the seed within, provi- 
ded with its own proper integuments. 
The name has been restricted to the seed-like fruits of simple 
pistils, such as those of the Buttercup (Fig. 628,629), Anemone, 
Clematis, and Geum. The style in some species of the latter 
remains on the fruit as a long and feathery tail, in others as a 
short and hooked one, both being agents of dissemination. The 
grains of the strawberry (Fig. 653) are also akenes. ‘The name 
is extended to all one-celled seed-like fruits resulting from a 


FIG. 625. Samara or key of White Ash, Fraxinus Americana. 626. That of White 
Elm. Ulmus Americana. 627. Double samara of Red Maple, Acer rubrum. 

FIG. 628. Achenium of a common Buttercup. 629. Vertical section, showing the 
seed within. 


ITS KINDS. 295 


compound ovary, and even when invested with an adnate calyx- 
tube. Of the latter is the fruit of Composite. (Fig. 630-635.) 
Here the tube of the calyx is incorpo- 

rated with the surface of the ovary ; 

and its limb or border, obsolete in some \y4/ |) /\/4jy 
cases (Fig. 630), in others appears Wh ily 
as a crown or cup (Fig. 651), or set of Man i 


\ Nh 
teeth or of scales (Fig. 652, 633), or as \ | 
a tuft of bristles or hairs (Fig. 634, \ 


634 


632 


635), &e., called the Pappus. In the Lettuce and Dandelion 
(Fig. 635), the achenium is rastrate, or beaked, 7. e. its summit 
is extended into a slender beak. An akene with adnate calyx 
has been termed a Cypse.a. 

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. me 

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 haying disappeared during its 
growth (536) ; as in the Hazel, 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 seales). 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 Cupule; such as the cup at 
the base of the acorn, the bur of the chestnut, and the leaf- 
like covering of the hazel-nut. The name Glans 
(sometimes Gland in English) is technically applied 
to such nuts, this being their classical Latin name. 

563. The fruit of the Walnuts and Hickory is 
apparently a kind of drupaceous nut, or something 
intermediate between a stone fruit and a nut. But 
certain monstrosities give reason for supposing that 
the seeming exocarp (041), which in Hickory 
hardens and at maturity dehisces*in four valves, 
is of the nature of an adnate involucre. The cocoanut is a sort 
of fibro-drupaceous nut. 

569. Nutlet, or in Latin form Nucute (Nucula), is sometimes 
superfluously employed in a literal sense, as a diminutive nut.? 
Of late it has acquired a good and fairly legitimate use as the 
name of the seed-like, or rather akene-like, closed parts or lobes, 
of crustaceous or other hard texture, into which certain bilocular 
or plurilocular pericarps separate at maturity, 7. e. for the seg- 
ments of a schizocarp, 571, which resemble akenes.? These are 
sometimes carpels, sometimes half-carpels, as in Verbena, also 
in Borraginaceee and Labiatee (in which the segments are greatly 
separated in the ovary), and sometimes, as in Nolana, they are 
portions of compounded carpels which have been exceedingly 
multiplied by chorisis. 

570. There are complete transitions between dry nutlets, with 
a thin and herbaceous epicarp, and the pyrene (574) or stony 
inner portion of such carpels when drupaceous or composing a 
drupe of two or more stones. It is therefore a hardly incongru- 
ous and very convenient use which extends the term nutlet to 
include these small seed-like stones also, as, for example, to 
those of Holly, Bearberry, Hawthorn, and the like. 

71. The pair of achenium-like or often samara-like carpels, 


1 Nut and akene, between which there is no fixed distinction, will cover 
this ground. The fruit of Cyperaceex, for instance, is truly an achenium, 
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, Beeckler, a caryopsis. 

2 Cocci (sing. Coccus, from a Greek word for kernel) is another name for 
fruit-carpels, or separating lobes of a dry pericarp, as well for dehiscent ones 
(of Euphorbia) as for indehiscent. Hence such lobed or partible fruits 
are said to be dicoccous, tricoccous, &e., according to the number of lobes or 
carpels. 

FIG. 638. Acorn (nut) of White Oak, with its cup. or enpule. 


2- 


ITS KINDS. 297 


united by their inner face but separating entire at maturity, 
which constitute the fruit of Umbellifersze, takes the name of 
Cremocarp (Lat. Cremocarpium); and the halves are called 
Mericarps. 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 
earpel. 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 Scuizocare. 
The component carpels of such a fruit were long ago named Car- 
cerules (carceruli, little prisons) by 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 

075. 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 
the ripening of which the 
outer portion of the pericarp 
becomes fleshy or pulpy, and 
the inner stony or crustace- 
ous, 7. é. divides into sarco- 
carp and putamen. (541.) 
But the name is extended to oe at 
pericarps of similar texture resulting 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, Rhamnus, &c. The several 
pericarps of the aggregate blackberry and raspberry are diminu- 
tive drupes or DruPe.ets. 

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 by a seed.1 
Bearberries (Arctostaphylos) and Huckleberries (Gaylussacia) 
are good illustrations of this. The seed-like endocarps of this 


1 The term Ac/inus, the original name of such a berry as a grape, has been 
used in descriptive botany for a small drupe or drupelet, and the ripened 
carpels of Rubus have been termed acini or acines, but without discriminating 
them from berries. 

FIG. 639. Vertical 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 PrrEN&; and the fruits are dipyrenous, tripyrenous, 
tetrapyrenous, &c., according as they contain two, three, or four 
pyrene. When the sarcocarp is thin and dries up at maturity, 
these pyrenz pass by gradations into nuculz (569) or nutlets: 
hence pyrene 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), of 
parchment-like or (in Hawthorns) bony texture, 
enclosed in flesh which morphologically belongs 
to adnate calyx and receptacle; as may be 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 (395) ; in the 
apple and pear, the inner or core-portion of the 
flesh is of the nature of disk, investing the carpels. 
In the fruit of Hawthorns, the carpels become bony 
pyrene (074), and so the fruit is drupaceous, is 
indeed nothing more than a syncarpous drupe. 
In Eriobotrya, or Cumquat, the carpels becoming 
very thin and membranaceous, the pomaceous 
fruit is in fact a kind of berry. 

576. The Pepo, 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 Cucurbitacez, 
fleshy internally and with a hard 
or firm rind, all or part of which 
is referable to the adnate calyx 
completely incorporate with the 
ovary. This is either one-celled 
with three broad and revolute. 
parietal placente, or these pla- 
cent, borne on thin dissepiments, 
meet in the axis, enlarge, and 
spread, unite with their fellows 

‘ on each side, and are reflected to 
the walls of the pericarp, next which they bear their ovules. As 
the fruit enkarges, the seed- -bearing placentse usually cohere 
with the walls, and the partitions are obliterated, giving the 


FIG. 641. Pome or apple in transverse section. 642. Quince in vertical section: the 
inner flesh answering to disk in the apple and pear is here wanting. 

FIG, 643. Section of the ovary of the Gourd. 644. Diagram of one of iis constituent 
earpels. 


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 placente. 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 (lig. 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 C7 
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 may need to be 
taken into account. The best name for it is 
that of Syxcarprom, or in English 
form Syncarp. But the term has 
been applied to multiple fruits as 
well.! In Hydrastis, the numerous | 
earpels imbricated on the upper 
part of the torus are baccate, that 
is, become berries ; in araspberry, 6i7 
the seemingly baccate grains are drupaceous (being drupelets, 573), 


1 The syncarp which is a gynecium might be designated a simple syn- 
carpium ; that which is an inflorescence, a complex syncarpium, which may be 
biflorous, pauciflorous, or multiforous. 

FIG. 645. The larger Cranberry, Vaccinium (Oxycoccus) macrocarpon; the one 
transversely divided. 

FIG. 646. Vertical section of half of a blackberry (of Rubus villosus), enlarged; and, 
647, of one of its drupelets more magnified. 


300 THE FRUIT. 


and, slightly cohering together (though without organic union), 
they fall as one body from the conical dry torus at maturity. It 
is the same in blackberries or bramble-berries (Fig. 646, 647), 
except that the drupelets persist on the torus, which partakes of 
the juiciness. In the aggregate fruit of Magnolia (Fig. 648-650), 
such carpeis, imbricated over one another, cohere more or less 
at all contiguous parts, and 
become drupaceous ; neyer- 
theless, at maturity each 
opens dorsally, allowing the 
seeds to fall out: in age it 
dries and hardens, and also 
separates from its conneec- 
tions, and so_be- 
comes a follicle, but 
with the remark- 
able peculiarity of 
dorsal instead of 
ventral dehiscence. 
(Fig. 650.) 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 Anthocarpous Fruits are those of which sq@me 
conspicuous portion of the fructification neither belorigs to the 
pistil nor is organically) united 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 
Anthocarp or Anthocarpium. This condition may occur either 
in simple, in aggregate, or in multiple fruits. 


1 The aggregate fruit like that of Rubus (named by some Conocarpium, 
by others an -Lterio, Erythrostomum, &c.) was termed by Dumortier a Drupe- 
tum. <A similar aggregation of baccate carpels he termed a Baccetum ; of 
follicles, a Follicetum, &¢. All such names may look well ina system; but 
they are both superfluous and unmanageable in phytography. 


FIG. 648. Aggregate fruit of Umbrella-tree, Magnolia Umbrella, reduced in size; a 
seed from a lower dehiscent carpel hangs on « thread. consisting of a tuft of extensile 
spiral ducts unravelled. 649. Same in longitudinal section. 650. One of the carpels 
detached, at full maturity, dried up, dorsally dehiscent, exposing the pair of seeds of 
the natural size. : 


ITS KINDS. 301 


5381. Gaultheria procumbens, 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. 645. 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 achenium 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 (889, Fig. 406, 655), the akenes or 
true fruits dispersed over the surface being 
apparently insignificant. Equally in many 
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 pinc-apple. 

583. Multiple or Collective Fruits + are those which result from . 
the aggregation of several flowers into one mass. The simplest 
of these are those of the Partridge-Berry (Mitchella, Fig. 467), 


bol 


65383 


1 Collective is the preferable name. The term mu/tiple was applied by 
DeCandolle to what are here (following Lindley) called aggregate fruits ; 
‘and the aggregate fruits of DeCandolle are here called multiple or collective. 
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 procumbeps, with enlarging calyx partly 
covering it. 652. Same, more advanced, ani 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 
are, in fact, dense forms of inflorescence, with the fruits or floral 
envelopes matted together or coherent with each other; and all 
or some of the parts succulent. The grains of the mulberry 
(Fig. 654-656) are not the ovaries of a singie flower, like those 


659 655 656 


of the blackberry, which it superficially resembles: they belong 
to as many separate flowers ; 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 Strawbgrry Blite may 
resemble strawberries ; but the pulpy part is the Calyx of many 
flowers, not the succulent receptacle of one. In the pine-apple, 
the flowers are spicate or capitate on a simple axis, which grows 
on beyond them into leafy stem; this when rooted as a cutting 


3. Fruit compound (ovaria compound), Syncarpr1; 4. Collective fruits, 
Anthocarpi. . 

Later, in his “ Elements of Botany,” Lindley reduced the classes to 
two: 1. Simple Fruits, those proceeding from a single flower; 2. Multiple 
Jruits, those formed out of several flowers. 


FIG. 654. A mulberry. young. 655. One of the fleshy grains at flowering time, show- 
ing it to be a pistillate blossom with fleshy calyx. 656. Thesame later, with the succu- 
lent sepals in transverse section. 

FIG. 657. A young fig. 658. Longitudinal section of the same later, but in flowering 
time. 659. 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 juicy 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 familar 
either in ordinary language or in descriptive botany. ; 
584. The Syconium or Hypanthodium, the Fig fruit. (ig. 657— 
659.) This results from a multitude of flowers concealed in a 
hollow flower-stalk, if it may be so called, which becomes pulpy 
and edible when ripe; and thus the fruct 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 
Mito very 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 by. 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 small 
orifice which is found in the fir. ; 
_ 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 beiongs to the 
Pine or Fir cone, the peculiar fruit of 
Coniferze (507), in which naked seeds 
are borne on the upper face of each 
fructiferous scale (Fig. 661), or gome- 
times in their axils. 
Such a cone when 
spherical, and of 
thickened scales 
with narrow base, as 
that of Cypresses, 
has been termed a 
GALBULUs, an unnecessary 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. 

586. 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 by one (chiefly the ventral) suture, . . . . . . . FOLLicte. 
Opening by both sutures;;.-: 2 2. 2 2°90) & 2°) See 
Or transversély jointed; 2) 22. 1 3) 2 
Dry and dehiscent, bi-pluri-carpellary,. . . . . + . . . « CAPSULE. 
When its dehiscence is circumscissile, . . .- oJ. «= gJPyoge 
When dehiscent by two valves from two eel een. . SILIQUE. 
A short and broad silique,. . . : 3 32 SEECrEE 
Dry and bi-pluri-carpellary, splitting = Gneseedea carpal . SCHIZOCARP. 
The dimerous schizocarp of Umbellifere, . . . . . . .CREMOCARP. 
Each of its halves or carpels,. . - . . . Hemicarp or MERICARP. 
The akenelike or nut-like Pe into which Schizocarps generally 
Mivides sa.) 0 . . Nucures or NUTLETS. 
Dry and indehiscent, queeetet one swordceded! 
Winged} 2.04 eer MSS OMe hfe os See EL 
Wingless, and with the 
Thin pericarp consolidated with the seed, . . . . . . Caryopsis 
Thin pericarp loose and not filled by the seed, . . . . . Urricie. 


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 cupule orinvolucre, ... ... . . . GLANS. 


Fleshy and indehiscent, 

Heterogeneous in texture, having 
A stone (putamen) 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 Gourd Family), . . . Prpo. 
From a superior ovary (confined to Orange Family), Hrsperrpicum. 
Homogeneous, fleshy throughout, .... . . . + + « «+ BERRY. 


THE SEED. 305 


CHAPTER VII. 
THE SEED. 


507. Tue Seep is the fertilized ovule (515), with embryo 
formeG 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. 665. 

588. The SrEep-staLK or PoposPERM, when there is one, is 
the funiculus of the ovule (516), and retains this name. So 
also do the Cuaraza, Ruaprue, and Hitum; the latter being the 
scar left by the separation of the seed from a 
its funiculus or directly from the placenta. 
The foramen of the ovule, now closed, is 
the MicropyLe 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 crustaceous in texture, 
it takes the name of Trsra, which is equivalent to seed-shell. 
It has also been named Spermoperm (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 hilum (a); testa (5); tegmen (c); albumen (4); 
embryo (¢). 664. Vertical section of the orthotropous seed of Helianthemum Cana- 
dense, with its funiculus, a. 


306 THE SEED. 


the nucleus, or loose and cellular (as in Pyrola-seeds), or vari- 
ously appendaged. 

591. The inner coat, ~1lled TreGmMeN and sometimes Enpo- 
PLEURA, 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. Iwo 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 
Epilobium, and at both ends in several 
Apocynaceze. In the Cotton-plant, very 
long and soft hairs, admirably adapted for 
spinning, thickly cover the whole seed- 

65 = 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 (563, 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 at a propitious time or place. In many but not all 
Polemoniacez (notably in Collomia), in certain Acanthacee, 
such as Ruellia tuberosa (and equally in certain Composite of 
the Senecio tribe and in Salvias, &c., among Labiatee, 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- 


FIG. 665. Winged seed of Trumpet Creeper, Tecoma radicans. 666 That of Catalpa, 
becoming comose: the body divided lengtliwise through the embryo. 
FIG. 667. Comose sced of Milkweed, Asclepias Cornuti. 


ITS COATS AND APPENDAGES. 307 


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 sma i 
and light seeds to the soil in time of 
rain, or to moist ground, favorable to 
germination. In cress and flax-seed, i 
the abundant mucilage developed when a——)- 
wetted comes from the gelatination of 
epidermal cell-walls, and 
subserves a similar use. 
.094. While the testa in 
many seeds is hard and 
crustaceous or bony, imitat- 668 64) 
ing the pericarp of a nut, in others (such as Peonia) it becomes 
berry-like (baccate), and in Magnolia, drupaceous.’ (Fig. 668- 
671.) These may also be regarded as adaptations for dissemi- 


67 


nation, here by 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. 106, from which the accompanying figures and Tig. 
589-597 are reproduced. 


FIG. 668. Forming seed (one eighth of an inch long) of Magnolia Umbrella; the 
rhaphe toward the eye. 669. Magnified view of the same divided lengthwise through 
the rhaphe; the outer coat, a, beginning to form a hard inner layer, a’. Within and 
distinct from this is the inner coat (b), 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. 

FiG. 670. A nearly full-grown seed, of the natural size. 671. Longitudinal section, 
emiargen, suowing the crustaceous or stony inner stratum of the testa well developed; 
the parts lettered as in Fig. 669. 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 and imbedded in the fleshy drupaceous 
testa, as shown in Fig. 670-672. 

596. Crest-like or other appendages are not uncommon either 
on the rhaphe or at the hilum. These are outgrowths produced 
during the development of the ovule into the seed. In Sangui- 
naria, such a crest develops from the whole length of the rhaphe 


672 675 674 65 


(Fig. 673) : in Dicentra, Corydalis (Fig. 674), &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, Ricinus (Fig. 675), &e., is produced by an out- 
growth of the external orifice of the ovule, the micropyle of the 
seed. This properly takes the name of Caruncie. But the 
two terms are not always discriminated. By further develop- 
ment, either of these may give rise, in certain seeds, to an acces- 
sory covering called 

097. The Aril or Arillus. This term, rather vaguely employed 
by Linnzeus, was first well defined by Geertner. The true arillus 
is an accessory seed-covering, more or less incom- 
plete, formed between the time of fertilization and 
the ripening of the seed, by a growth from the apex 
of the funiculus (when there is any) at or just be- 
low the hilum, in a manner similar to that in which 
the coat or coats of the ovule are formed. That 
of Nymphiea (Fig. 676) is a typical example ; only 
the arillus is developed from the funiculus at a point distinctly 
below its apex: here a ring forms, which grows into a cup, and 
this is soon extended into a sac, loosely enclosing the seed, and 
open at the top. This is membranaceous : commonly it is fleshy. 
When there is absolutely no funiculus. the aril may originate 
from the placenta, as it does in Podophyllum, in which most of 


FIG. 672. Anatropous seed of Sarracenia purpurea, with very salient rhaphe. 673. 
Same of Sanguinaria or Bloodroot, with rhaphe crested for its whole length 674. Seed 
of Corydalis aurea, with crest or strophiole, attached at or near the hilum. 675. Seed 
(suspended) of Ricinus, with its caruncle. 

FIG. 676. Seed of White Water-Lily, Nymphaa odorata, in its loose and thin arillus. 


~) 


ARILLUS, ALBUMEN. 300 


S, 


the pulp of the berry consists of these fleshy arils, much com- 
pacted. (Fig. 677, 678.) 

598. The laciniate aril of the nutmeg (mace) and, it is said, 
the bright red and pulpy 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 deyeloped an 
aril of this sort, it would 677 673 
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 (Arillodium) 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, &e.), 
is the name generally employed by systematic 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 are 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, &e. 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 may 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 may disappear 
altogether, as in the seeds of Maple, Almond, Squash, Pea, and 
the like, which are therefore said to be exalbuminous. 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 peltatum; the pulp of 
the latter mainly of the nature of arillus. investing the seeds. 678. The arillus of ong 
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 (25, note); but neither name has in 
systematic botany displaced the earlier one of Grew and Geert- 
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 ENpospErmM, and that 
formed without, which takes the name of PEerisperM. ‘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. 

603. In most seeds the albumen is endosperm: in Canna it 
is all perisperm. In Nymphvea and its allies (except Nelum- 
bium, which has none) mostof it is perisperm ; 
but a thin and condensed layer of endosperm 
surrounds the embryo, 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 quan- 

679 tity of endosperm or inner albumen., 

604. When the nucleus of a ripe seed is hollow, as in the 
cocoanut and nux vomica, the formation of endosperm, which 
usually begins next the wall of the embryo-sac, has not proceeded 
so as to fill the cavity. The embryo-sac in the cocoanut attains 
enormous size, and the cavity is filled by the milky fluid. 

605. The texture or consistence of the albumen differs greatly. 
It is fartnaceous or mealy when, consisting mainly of stareh- 
grains, it may readily be broken down into a powder, 
as in wheat, buckwheat, &c. ; o¢/y, when saturated 
with a fixed oil, as in poppy-seed; fleshy, when 
more compact, but readily cut with a knife, as in 
the seed of Barberry ; muctlaginous, when soft and 
somewhat pulpy, as in Morning Glory and Mallow, 
but when dry it becomes fleshy or harder ; corneous, 

6S) when of the texture of horn, as in coffee and the 
seed of Caulophyllum ; and even ony, as in the vegetable ivory, 
the seed of Phytelephas. It is mostly uniform ; but in the nutmeg, 


FIG. 679. Longitudinal magnified section of a seed of Black Pepper; showing the 
large episperm, the small endosperm in the persistent embryo-sac, and in this the 
minute embryo. 

FIG. 680. Longitudinal section of a seed of the so-called Papaw, Asimina triloba, 
with ruminated albumen and minute embryo. 


THE EMBRYO. Sit 


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 ruménated. 


632 Got 


606. The Embryo,’ 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 cotyledons) 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 (Considérations sur les Corps organisées), in 1762, and was introduced 
into systematic botany at about the same time (1763) by Adanson: it was 
taken up by Gertner in 1788. Jussieu in the Genera Plantarum (1789) held 
to the term Corculum (the cor seminis) which came down from Cesalpinus. 

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 (anatropous), enlarged; with hilum or scar (a), rhaphe 
(6), and chalaza (ce) indicated. 682. Vertical section of the same, showing the straight 
embryo in the axis of the mealy albumen. 

FIG. 683. Vertical section of the (orthotropous) seed of Buckwheat, showing the 
embryo folded round in the mealy albumen. 

FIG. 684. 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 
ce. the chalaza, d, the testa, e, the tegmen,.f, 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. 


312 THE SEED. 


608. The opposite extreme is an embryo (as in Fig. 686) 
which appears as a mere speck in the albumen, but 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) of a dicotyledonous embryo, indicating 
the future cotyledons. Indeed, in Monotropex, Orobanchacee, 
and some other parasitic dicotyledonous plants, and in Orchids 
among the monocotyledonous, the embryo 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 
radicular 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 
Helianthemum (Fig. 664) and Pepper (Fig. 679), the radicle 
necessarily points directly away from the hilum.’ 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 different way, 
in the campylotropous seed (Fig. 689, 
690); while in the amphitropous the 
radicle points away from the hilum later- 
ally. 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 (caulicle), or rather its relation to the hilum, are 

Antitropous, when the embryo directs its radicle away from the hilum, as 
it must in all orthotropous seeds ; 

Orthotropous, also homotropous, when directed to the hilum (more strictly to 
the micropyle close to the hilum), as in anatropous seeds. These two terms 
are still employed by many botanists, although superfluous when the ovule 
or seed is stated to be anatropous or orthotropous, &c. And the term 
orthotropous, so used, is liable to be confused with orthotropous as applied 
to the ovule. 

Richard, moreover, termed the embryo amphitropous when curved or coiled, 
as in Chickweed (Fig. 689) and all such campylotropeus seeds; and hetero- 
tropous 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. 689. Campylotropous seed of common Chickweed, magnified. 690. Section ot 
the same, showing the embryo coiled into a ring around the albumen. 


THE EMBRYO. 313 


of the embryo within, and of its parts, may often be inferred 
withont 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 systematic writers; who employ 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 peripherie. 

613. The embryo may be very variously folded or coiled in 
the seed. The two cotyledons, instead of plane and straight, 
may be crumpled; or they may be simply convolute or rolled up 
from one edge, as in Calycanthus (Fig. 691) ; or etreinately con- 


GOL 692 695 


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 Cruciferze as to need special reference. Namely, when 
cotyledons 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 Cruciferze by 


FIG. 691. Convolute embryo of Calycanthus, the upper half cut away. 

FIG. 692. Seed of a Cruciferous plant (Sisymbrium), with incumbent cotyledons, 
divided. 693. Embryo of the same 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, 7. e. leaves at the 
first nodes alternate (39); and 

Dicotyledonous, with a pair of cotyledons, 7.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 Conifer (Fig. 48, 49), occasionally and abnormally in sundry 
ordinary dicotyledonous species. 

615. There are several embryos of the cotyledonous type in 
which one cotyledon is smaller than the other, viz. the inner 
one when the embryo 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 embryo 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 were 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. Tue PrivcipLes or 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 History, 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 study. 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 came from common 
parents or near ancestors. 

620. The recognition of the perennial succession of similar 
individuals gives the idea of Spectres. The recognition of un- 
equal degrees of likeness among the species is the foundation 
of GreneRA, Orpers, 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, arid 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 things; but, although the tend- 
ency to individuation begins with life itself, it is completely 
realized only in the higher animals. 

622. 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, 156.) 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. 

623. 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 off- 
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 and 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. 

624. 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. 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 are 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 being 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 Abhandl. Akad. 
Wissenschaften zu Berlin, 1853), Das Individuum der Pflanze, &c., and a 
translation by C. F. Stone in Amer. Jour. Sci. ser. 2, xix. xx. 1855. 


THE PRINCIPLES OF CLASSIFICATION. 317 


cisely similar grounds, carry the analysis a step farther, and 
regard each phytomer (16) 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 cryptogamous 
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.t Bnt, 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 ox 
beings which have come from one stock, in virtue of that mosé 
general fact that likeness is transmitted from parent to progeny. 
Among the many deiinitions, that of A. L. Jussieu is one of the 
briefest and best, since it expresses the fundamental conception 
of a species, 7. 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 relationslip 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 Jast 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 progeny may come to differ) 
strikingly in some particulars. So two or more forms which 
would have been regarded as wholly distinct are 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 genera- 
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. 

628. 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 
preserved, led on to further development, and relatively fixed. 

629. 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 that spring from them will show marks 
of this different treatment in their appearance. Such differences 
are 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 
spontaneous 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 the 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, &e. 

630. Some varieties of cultivation originate in comparatively 
slight deviations from the type, and are 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.”1 That is, some seedlings, or some shoots, are 
unlike the rest in certain particulars.” 

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.* 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, z.e. the tendency to inherit from grand- 
parents, great-grand-parents, &c. Atavisin, 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 arc— must be 
much repressed by the prevalent cross-fertilization which takes 
place among tke 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, Rye, Cabbage, Lettuce, Radishes, Peas, &c., and 


1 Both the technical English term. Sport, and its Latin equivalent, Lusus, 
are sometimes used for bud-yariation 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 sced-bed does not suggest this view. Negeli, 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 TAXONOMY. 


indeed of nearly all our varieties of cultivated annual and biennial 
esculent plants, as well as of several perennials, many of which 
have been fixed through centuries of domestication, while others 
are of recent establishment. What is now taking place with 
the Peach in this country may convince us that heritable varieties 
may be 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 years than 
in shrubs or trees. Varieties of this fixity of character are called 

633. Races (Lat. Proles). 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.’ In most of these, the position of species and variety is 
more or less arbitrary or accidental, and capable of interchange. 
What is called the species may be only a commoner or better- 
known form, or the one first recognized and named by botanists ; 
whence the other forms as they come to be recognized are 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 type. 

634. 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 are sometimes called Subspecies. 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; or 
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.* 


! The Horseradish and a few other plants of spontaneous growth, which 
through long dependence on bud-propagation seem to have lost the power 
of setting seed, can hardly be called varieties. 

2 Darwiniana, 35. 

8 Wherefore, since we hardly need the term race in the restricted sense 
of seed-propagated variety, it is sometimes convenient to use it in the man- 
ner proposed by Bentham (Anniversary Address to the Linnean Society, 


THE PRINCIPLES OF CLASSIFICATION. $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.’ In nature, cross-breeding doubtless re- 
presses variation or prevents the segregation of varicties 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 by 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 very similar, refusing to unite. But, on the whole, 
there seems to be few nearly 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 

658. 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 may 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, Subspecies 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 Miztus ; French, Métis. 
2] 


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 are 
fully fertile per se, divide in the offspring, partly in the first 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 adventive embryo-formation, 533.) 

639. 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 species 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 there 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 


! According to Naudin in Comptes Rendus, xlix. 1859, & lv. 1862. See 
also Naudin’s memoir on hybridity in plants in Ann. Sci. Nat. ser. 4, xix. 
1863, pp. 180-205, & in Mem. Acad. Sci. . . . For the literature on vegetable 
hybrids, see Keelreuter, Nachricht, &c., 1761, and Appendices, 1763-1765; 
Herbert, on Amaryllidacee, 1837 ; C. F. Gertner, Versuche und Beobachtun- 
gen ueber die Bastarderzeugung in Pflanzenreich, 1849; Wichura, Die 
Bastardbefruchtung im Pflanzenreich, erliutertert an den Bastarden der 
Weiden ; and the memoir of Naudin referred to. 


THE PRINCIPLES OF CLASSIFICATION. 323 


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. Linnzeus 
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 Class ; 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 
are 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 be 
equivalent terms. If only 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 Cat 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 Jackal, &e. The 
languages of the most barbarous as well as of civilized 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. 

643. 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 may 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, the pinnatifid or sinu- 
ate leaf might have been thought as essential to the Oak 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 
European and American Oaks; but in numerous Asiatic species 
the cup bears concentric or spiral lamellze instead, and in others 
the cup takes the form of a naked and closed sac. Maples have 
palmately-veined 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. Sy Ay 


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. Famity in botany is synony- 
mous with order: at least natural orders and families (however 
distinguished in zoology) have always in botany been inter- 
changeable terms, and will probably so continue.! 

646. As examples of orders in the vegetable kingdom take 
the Oak family, composed of Oaks, Chestnuts, Beeches, &e. ; 
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. Kingdom 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., may 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. 

649. The sequence of groups, rising from particular to univer- 
sal, is Species, Genus, Order, Class, Kingdom; or, in descending 
from the universal to the particular, 


Kinepom, 
Crass, 
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. 


650. This is the common framework of natural history classi- 
fication. All plants and all animals belong to some species ; 
every species to some genus; every genus to some order or 
family ; every order to some class; every class to one or the 
other kingdom.t But this framework, although all that is re- 
quisite in some parts of natural history, does not express all the 
observable gradations of relationship among species. And even 
gradations below species have sometimes to be classified. The 
series 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. 

651. 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? (or sub- 
kingdom), the Phzenogamous and the Cryptogamous. 

652. The grades intercalated into the long-established sequence 
of Class, Order, Genus, and Species, with new names, are mainly 
two, 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 Botany, 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. 

655. 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 Not recognizing Heckel’s third kingdom of Protista, consisting of those 
lowest forms of being from which the animal and vegetable kingdoms 
emerge. 

2 Answering to the French Embranchement in zoology. For this it is pro- 
posed to use the word Dyirision (Divisio): see Laws of Botanical Nomencla- 
ture adopted by the International Botanical Congress held at Paris in 
August, 1867 ; together with a Historical Introduction anda Commentary, by 
Alph. DeCandolle, — English translation, London, 1868 ; the original French 
edition, Paris, 1867. Perhaps no better name can be found; but the elder 
DeCandolle brought Divisio into common use for a grade subordinate to 
tribe. Endlicher employed the term Regio. We have used Series, and 
much prefer it. 


THE PRINCIPLES OF CLASSIFICATION. 327 


example, if Dicotyledones and Monocotyledones be the two 
classes of Phznogamia, the former (and only the former) is 
divided upon very important characters into two branches, of far 
higher rank than the cohorts, viz. the Angiosperme and the 
Gymnosperme, which take the name of Sunctasses. 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 Suprrises. Genera may comprise sections of 
species which might almost as well rank as genera themselves : 
to mark their importance and pretension (which may come to 
be allowed), they are termed SusGenera. Finally, forms which 
are ranked as varieties, but which may establish a claim to be 
distinct species, are sometimes termed Scupspecres. 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 
always in the following sequence : — 

KINGDOM, 
Series or Division, or Sub-kingdom, 
Grass: 
Subclass, 
Cohort, 
OrD_ER or Family, 
Suborder, 
TRIBE, 
Subtribe, 
GENUS, 
Subgenus, 
Section, 
Subsection, 
SPECIES, 
Subspecies or Race, 
Variety, 
Subvariety. 


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, however graye, are sup- 
posed to have arisen from subsequent variation, and the more 
marked differences to have become fixed through heredity. This 
is included in the idea of species. Descent from a common 
origin explains the likeness, and is the only explanation of it. 

658. But what is the explanation of the likeness between the 
species themselves? As respects nearly related species, the answer 
is clear. Except for practical purposes and in an arbitrary 
way, no certain and unfailing distinction can be drawn between 
varieties of the highest grade and species of closest resemblance. 
It 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. 

659. 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 the 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 which we recognize as species. But 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. Just this is accounted for by 

660. 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 rigidly destroying all plants which do not show some desirable 
variation, breeding in and in from these, with strict selection of 


THE PRINCIPLES OF CLASSIFICATION. 3829 


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.! 

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.” 


350 TAXONOMY. 


moisture or dryness (which are transient and comparatively 
unimportant), variation, or the unlikeness of progeny to parent, 
is occult and inexplicable. If sometimes called out by the 
external conditions, it is by way of internal response to them. 
In Darwin’s conception, variation of itself does not tend in any 
one particular direction: he appears to attribute all adaptation 
to the sorting which results from the struggle for existence and 
the survival of the fittest. We have supposed, and Negeli takes 
a similar view,’ that each plant has an internal tendency or pre- 
disposition to vary in some directions rather than others; from 
which, under natural selection, the actual differentiations and 
adaptations have proceeded. Under 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 series of 
questions which without it are totally unanswerable. It is sup. 
ported by vegetable paleontology, 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 are impressed upon their 
structure as well as traceable in their geographical and paleon- 
tological 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 of small if any importance to the plant’s well- 
being, and that they run like threads through a series of species 
of the greatest diversity in habit, mode of life, and particular 
adaptations to conditions.’ 


1 Entstehung und Begriff der naturhistorischen Art. Zweite Auflage, 1865. 
2 This is a corollary of natural selection, which can take effect only 
upon useful characters, 7. ¢. upon structures which play some 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 
speciiic modification. Such modifications are too slow to effect 
in any wise the stability and practical application of botanical 
classification. 


Section II. BorantcaLt CLASssIFICATION. 


663. Natural and Artificial Classifications may 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 diiferent 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 by Linnzeus (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 
naturai classification. 


in the life of the plant, and which therefore undergo modification under 
changing 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, Rubiacee are known by insignificant stipules, Ano- 
nace by ruminated albumen, Rhamnacee by a valvate calyx and stamens 
before the petals, &e. Paradoxicat as it may seem, it is not although, but 
because, they are of small biological importance that they are of high clas- 
sificatory (7. e. of genealogical) value. 

On considerations like these, characters are divided into adaptive 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, &e. ii 


332 TAXONOMY. 


664. No artificial classification of plants could fail to be 
natural in some portions and some respects; because plants 
whick agree in any point of structure likely to be 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 of plants, even 
if these were fully 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 
system and throughout its parts, has been the task of the leaders 
in botany from the beginning of the science until now; and the 
work is by no means completed. 

665. Linnzus was perhaps the first botanist to distinguish 
clearly between a natural and an artificial classification. He 
labored 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. 

666. Ante-Linnean Classification. Linnzeus, in his Philosophia 
Botanica, 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 
Linneus made a primary division of the vegetable kingdom into 
Trees and Herbs, referring the larger shrubs to the former and 
the under-shrubs to the latter,—an arrangement which began 
with Theophrastus and was continued by Ray and Tournefort. 

667. The three most important names in botanical taxonomy 
anterior to Linnzeus are those of Cesalpini, Ray, and Tournefort. 
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. 
Conrad Gesner of Zurich had somewhat earlier recognized this 
principle, but Cesalpini first applied it. 

668. A century later (1690-99) this principle was carried into 
practice by Rivinus (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. 

669. 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 cail natural orders or families, but 
which he unfortunately called genera. He noted the coincidence 
of nerved leaves with the monocotyledonous embryo, 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 
year 1694, 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 Linnzus 
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. Linnean Classification. Linnzeus, the great reformer of 
hotany 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- 
ficiai classifications (663), and deferring all endeavors to make 
the former available, except for genera, he devised a practical 
substitute for it, as a key to the genera, viz. his celebrated 

672. Sexual System, or arrangement of the genera under arti- 
ficiai classes and orders, founded upon the stamens and pistils. 
Although now out of use, this artificial classification has been so 
popular and influential, and has left so deep an impression upon 
the science and especially upon the language of botany, that it 
needs to be presented. The primary divisions are the classes, 
twenty-four in number. But the 24th class, Cryptogamia, con- 
sists of plants which have not stamens and pistils and conse- 
quently no proper flowers, and is therefore the counterpart of the 
remaining twenty-three classes, to which the corresponding name 
of Phanerogamia or, in shorter form, Pheenogamia (Phanogamous 
plants} has since been applied. These twenty-three classes are 
characterized by certain modifications and associations of the 
stamens, and have substantive names, of Greek derivation, ex- 
pressive of their character. The first eleven comprise all plants 
with perfect (7. e. hermaphrodite) flowers, 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 by names compounded of Greek numerals and the word 
andria (from «r7je), which is used metaphorically for stamen, as 
follows : — 


Class 1. Monanprta includes all such plants with one stamen to the flower; 
as in Hippuris. 

Dranpria, those with two stamens, as in the Lilac. 

TRIANDRIA, with three stamens, as in the Valerian and Iris. 

TETRANDRIA, with four stamens, as in the Scabious. 

PENTANDRIA, with five stamens, the most frequent case. 

HEXANDRIA, with six stamens, as in the Lily Family, &e. 

HeEpPTANDRIA, with seven stamens, as in Horsechestnut. 

OcranbrRIA, with eight stamens, as in Evening Primrose and Fuchsia. 

. ENNEANDRIA, with nine stamens, as in the Rhubarb. 

. DEcANDRIA, with ten stamens, as in Rhododendron and Kalmia. 

DopecaNnpDRIA, with twelve stamens, as in Asarum and the Migno- 
nette; extended also to include those with from thirteen to nine- 
teen stamens. 


Fee, SCOR TE OO ON ES CONS 


et 


673. The two succeeding classes include plants with perfect 
flowers having twenty or more unconnected stamens, which, in 


12. IcosanpRIA, are inserted on the calyx (perigynous), as in the Rose 
Family; and in 

13. Potyanprta, on the receptacle (hypogynous), as in the Buttercup, 
Anemone, &c. 


BOTANICAL CLASSIFICATION. ae 


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. Dipynamrta, including those with two long and two short stamens, 
as in the majority of flowers with bilabiate corolla. 


15. TerTRApYNAMr1a, 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. MonapELPuHIA (meaning a single fraternity), with the filaments 
united in a single set, tube, or column, as in the Mallow. 

17. DiapeLruia (two fraternities), with the filaments united in two 
sets or parcels, as in Corydalis and in many Leguminose. 

18. PotyapEeLpuia (many fraternities), with the filaments united in 
more than two sets or parcels, as in Hypericum. 

19. Syncenesia (from Greck 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 style : — 


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 
occupy separate blossoms : — 


21. Monecia (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. Diacia (two households), where they occupy separate flowers on 
different individuals ; as in the Willow, Poplar, Moonseed, &e. 

23. PotyGamia, 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 essentially flowerless ; or rather its 
organs of reproduction are more or less analogous to, but not 
homologous with, stamens and pistils. But, although Linnzeus 
suspected a sexuality in Ferns, Mosses. Algze, &c., there was no 
proof of it in his day. So he named the class, containing these, 


.24, CryproGAmiA, 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. 


336 


‘VINVOOLAANO 
‘VINVDATIOY 


*VIOWIC 
*VIOIDNO IAL 
*VINGNVNAY 


‘VIS 


UO NAG 


‘VIHATIAVAITOd 


VINATAOV ICT 
‘VIHA THOVNO[L 


“VINVNAGVULLAL, 


“VIN VNACIC 


‘VINGNVATOT 
“VIMGNVSOOT 
“VIMNAUNVOMCOCT 
“VINANVOCUC 
*VINGNVANNG 
“VINANVLOO 
“VINGNV LAT] 
“VINGNVXGIT 
“VINGNV ING 
VINAINVULAT, 
“VINGNVIWT, 
“VIMAINVICT 
“VINGNVNO|A 


sTeNpIA 
=IPUT JUSIAYIP O91} IO OM} IO ‘ours oy UL 
‘poyurudes 8101130 ‘yoaytod S1OMOTF Oo) JO ONLOS 
as SPEUPIATpUL JUdLOYIp UL 
* S[UNPLATpUL ous ot[} UL 


“saoyue aporyy Aq 
8}08 omy UvT} O10 UL syuUOUTETYy toy Aq 

S}OS OMY UL SPUOUULL TT Tot) Aq 
408 OLSUIS UV UL S}UOMIE]Y Afolyy Aq 


SUOTILYS JLOYS OMY PUB FuoyT AMoJ 
* SUOTUUS JALOT[S OA PUB SUOL OMY 


xA7TwO OT) 09 YUdAOYpY you foroNr 10 TZ 
* * xA[VO O17) 07 queLOT|pY ‘OMOUL 10 0% A 
. . . . . . . . . . 6b II 4 
; ; pen Ot: gape OL " 
. . . . . . . . . . . . . L “ce 
. . . . . . . . . . . . . ¢ “e 
. . . . . . . . . . . . . ¢ “ 
. (pan o* ik . pee . ps . . ¢ “ 
. . . . . . . . . . . . . 4 oe 


Foe fe.” sy fee ere! oteram ee Or ib 


I suoulNys 


‘SUSSVIO NVWNNI'T 


es 


OUHAL 


Sepia Al eer er oh Aly. 


auou 10 ‘poyvoouod s[ystd puv suouinys o1} 


*  g19MOTT 0}BrEdoS UT 


* Tys8td oY 07 JUaIOYpV suoUTRyS OT) } 


PY suoy 
jenboun jo 


| pur ‘10190 
youo IIA 


* TOTO TPHVO YQEM poyoouT0 


pojoouuooun 


Tysuoy 
wubo jo 


‘TOMO 
oles OY} UT 
‘ssid punoy y40q 
OL} WLOAy 
OPV 
SudUIeySs 
oy 


TO MUTA TVOLLCONAS 


“qsoyTue ur 
ststd puv 
suotUeys 


Suavy 
SINVIG 


BOTANICAL CLASSIFICATION. 337 


680. The orders, in the first thirteen classes of the Linnean 
artificial system, depend on the number of styles, or of the 
stigmas when the styles are wanting; and are named by Greek 
numerals prefixed to the word gynia, used metaphorically for 
_pistil, as follows : — 


Order 1. Monocynta, those with one style or sessile stigma to the flower. 
2. Dicyni4, those with two styles or sessile stigmas. 
3. TricyniA, those with three styles. 

4. TerracyniA, those with four styles. 
5. PentaGYNiA, those with five styles. 
6. HexaGynta, those with six styles. 
7. HeprTacynia, those with seven styles. 
8. OcroeyniA, those with eight styles. 
9. EnneacyntA, those with nine styles. 
10. DecaGynta, those with ten styles. 
11. Dopecacynta, those with eleven or twelve styles. 
12. Potyeynta, 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 sced-vessel or peri- 
carp, 7. e. the pericarp undivided. 


682. The 15th class, Tetradynamia, is also divided into 
two orders. which are distinguished merely by the form of the 
pod : — 


1. Srxricutosa; the fruit a silicle (561), or short pod. 
2. Siz1quosa; 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 Monanpria, with one stamen, Dianpria, with two 
stamens ; and so on. 

684. The orders of the 19th class, Syngenesia, are six, namely : 


1. PoLyGAMIA £QUALIS, 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 PoLyGAaMiA 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. Monocamta, where solitary flowers (that is, not united into a head) 
have united anthers, as in Lobelia. 


338 TAXONOMY. 


685. The 23d class, Polygamia, has three orders, two of them 
founded on the characters of the two preceding classes and 
bearing their names, and the third named upon the same prin- 
ciple, namely : — 

1. Mone@cra, where both separated and perfect flowers are found in 
the same plant. 
. Diacra, where they occupy two different plants. 


. Tri@cra, where one individual bears the perfect, another the stami- 
nate, and a third the pistillate flowers. 


ww bo 


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. Frrices, the Ferns. 

2. Musct, the Mosses. 

3. ALG#, which, as left by Linnzus, comprised the Hepatice, Lichens, 
&e., as well as the seaweeds. 

4. Fune1, Mushrooms, &e. 


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 Linnzeus, 
who pronounced it to be the first and last desideratum in syste- 
matic botany; and he early attempted to collocate most known 
genera under natural orders (e. g. Piperite, Pulme, Scitamine, 
Orchidee, Amentacea, &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 difficult problem was taken up by Linnzeus’s contemporary 
and correspondent, Bernard de Jussieu, who planted the botanic 
garden at Trianon with plants grouped into natural orders, but 
published nothing. His pupil, Adanson, who when a young 
man lived for several years in Senegal, and who was as remark- 
able for eccentricity as for erudition and ability, published in 
1763, in his Familles 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 Genera Plantarum secundum Ordines Natu- 
rales disposita, 1789, natural orders of plants, one hundred 


Pe ee ee ee Cle eee ee ee ae 


BOTANICAL CLASSIFICATION. 339 


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, 
Monocotyledones, and Dies yledones, adopted from Ray, with a 
change which was no improvement. For his Acotyledones, the 
Cryptogamia of Linnzeus, are the ‘+ plants without flowers” of 
Ray: they are, to be sure, destitute of cotyledons (though not in 
the manner of Cuscuta), because destitute of embryo altogether. 
The Acotyledones forming his first class, Jussieu divided the 
Monocotyledones into three classes upon single and artificial 
characters, namely upon the insertion of the stamens, whether 
hypogynous, perigynous, or epigynous ; and the Dicotyledones, 
into eleven classes on similar characters, preceded by a division 
into Apetale, Monopetale, Polypetale, and Diclines irregulares, i.e. 
first upon the character of the perianth, then upon the insertion 
of the stamens or in Monopetalee of the corolla. The following 
is the scheme : — 


Acotyledones. . . He See «ces CLASS OI: 
Stamens hypogynous . . . . . If. 
Monocotyledones perigynouses oe. 2. EE 
epipynousia as os. <= live 

Stamens epigyuouss e723) 0 127" V. 

Apetalous . PeLioynousy 4 Gece 30 ee VAL 

by pooynoust A ac oe, VE 

Corolla hypogynous . . . . .VIII. 

perigynous . Bore corel D.& 


epigynous: anthers connate X. 


epigynous: anthers separate XI. 


( 

Monopetalous 
Dicotyledones . 4 

( 


Stamens epigynous. . .. . . XII. 

Polypetalous . hypogynous . .. . XIII. 
perigynousi:, |. 8.52, 4 UNLV. 

Diclinous (also Apefalous) - . =. .- . . . . . 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 pheenogamous 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- 
gene and Endogene, appended to Dicotyledones and Monoco- 


340 TAXONOMY. 


tyledoneze, as it now appears should have been omitted. The 
giades of the Candollean system superior to the orders, in their 
final form, are mainly these : — 


pry. I. VASCULAR (more properly PHA2NOGAMOUS) PLANTS. 


Crass I. DicoryLeponowus or EXOGENOUS. 
Subclass I. THaLtamirLorous : petals /distinct) and stamens on the 
torus, 7. e. free. 
II. Catyciritorous: petals (distinct or coalescent) and 
stamens adnate to the calyx. 


III. CorotiirLorous: petals (mostly coalescent) not ad- 
nate to calyx, bearing the stamens. 


IV. Monocutamypeous: petals wanting. 


Crass IL. MonocotyLeponous or ENpoGENous. (No subclasses.) 


Diy. Il. CELLULAR (more properly CRYPTOGAMOUS) PLANTS. 
Crass I. ASTHEOGAMOUS: with sexual apparatus, and 
Vascular tissue. (Hquisetacew—Filices.) 
Only cellular tissue. (Musci and Hepatice. ) 


Cuiass II. AmpuiGamous: destitute of sexual organs and of other than 
cellular tissue. (Lichenes, Fungi, Alge.) 


691. Cryptogamous plants of all orders are now known to be 
provided with sexes; and the Jussixan divisions of the Dico- 
tyledones into Apetale (including Diclines), Monopetale, and 
Polypetale, 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. 

692. John Lindley in successive attempts (between 1830 and 
1845) variously modified, and in some few respects improved, 
the Candollean 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 effect 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 nixus (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 has no good Latin equivalent, and the 
term cohort (coors) 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 Coniferse and Cycadacez, 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 Angiosperme and the Gymnosperme. The 
latter has been elevated by the vegetable paleontologists 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 secunduin 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. Thallophyta, plants without proper axis of growth 
(developing upward as stem 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 A%theogamous Cryptogamia of 
DeCandolle, with which was wrongly associated a group of root- 
parasitic flowering plants (the Rhizanthez) which were fancied 
to bear spores instead of embryo in their seed; 2. Amphibrya, 
which answer to Monocotyledones ; and, 3. Aeramphibrya, which 
answer to Dicotyledons. These last contain five cohorts: 1. 
Gymnospermee ; 2. Apetale; 3. Gamopetale (the Monopetale 
of Jussieu better named): 4. Dialypetale (the Polypetale of 
Jussieu, &e.). 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, 7. 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 Thalamiflore with some of his Calyciflore) ; 
and under these the orders are arranged in cohorts, — fifteen 
cohorts in the Polypetale, and ten under three ‘‘ series ” in the 
Gamopetale. The remainder of this particular classification 
has not yet appeared in print, although partly sketched by its 
authors. It will generally be adopted in this country, with some 
occasional minor modifications. 

696. 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 flowers are reductions or 
degradations of polypetalous types, intercalates the Apetale or 
Monochlamydez among the Polypetalz. But this has never yet 
been done in a satisfactory manner, or without sundering orders 
which should stand in contiguity. 

697. 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 system, and especially 
in a lineal order, necessary as a lineal arrangement is, must 
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 In the first place, the relationships of any group cannot always be 
rightly 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 groups 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 classification. This might be expected from 
the nature of the case. For the naturalist’s groups, of whatever grade, are 
not realities, but ideas. Their consideration involves questions, not of things, 
between which absolute distinctions might be drawn, but of degrees of resem- 
blance, which may be expected to present infinite gradations. Besides, al- 
though the grades of affinity among species are 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 his works 
he is obliged to arrange the groups he recognizes in a lineal series; but 
each genus or order, &c., 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 Phanogams and leaving the 
lowest in contiguity with the higher Cryptogams. The Dicoty- 
ledons take precedence of the Monocotyledons in rank. Yeta 
part of them, the Gymnosperms, are much the lowest of all 
known Phenogams 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 tendency, 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 Gnetaceze and Loranthacez is 
not respected. (606, &c.) 

699. Nor can the angiospermous Dicotyledons be disposed 
lineally according to rank. The apetalous and achlamydecus 
must be the lowest. Some are evidently reduced forms of Poly- 
petale or even of Gamopetalze: the greater part cannot without 
violence be thrust into their ranks. The Gamopetal, 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 Composite 
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 
phznogamous natural orders, as another is to illustrate the 


344 TAXONOMY. 


cryptogamous orders. A synoptical view of the great divi- 
sions only, as at present received and named, is appended. 
Definitions and characters may be sought in the present and 
preceding chapters. 


SERIES I. PHAENOGAMOUS OR FLOWERING PLANTS. 


Crass L DICOTYLEDONS. 


Supcrass I. ANGIOSPERMS. 
Div. 1. POLtyPpetTaLous. 
Div. 2. GAmMopetTatous. (Monopetalous.) 
Div 3. APETALOUS. 


Suscrass Il. GYMNOSPERMS. 
Crass Il. MONOCOTYLEDONS. 
Div. 1. Spapicrovs. 


Div. 2. PETALOIDEOUS. 
Div. 3. GLUMACEOUS. 


SERIES II. CRYPTOGAMOUS OR FLOWERLESS PLANTS. 


Crass III. ACROGENS. 
Div 1. Vascurar. (Ferns and their allies.) 
Div. 2. CELLULAR. (Mosses and Liverworts.) 


Crass IV. THALLOGENS OR THALLOPHYTES. 


PHYTOGRAPHY. 345 


CHAPTER X. 
PHYTOGRAPHY. 


Section I. NOMENCLATURE. 


701. Payrocrapny 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 mostly European, their scientific names 
were mainly adopted from the ancient Romans or, through Latin 
literature, from the Greeks. Ex. Quercus, Prunus, Rosa, Rubus, 
Trifolium ; 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, 
&e. 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 7hea and 
Coffea. Of American aboriginal names, Hura, Guaiacum, and 
Yucea are examples. Some ancient names of plants commem- 
orated distinguished men. Ex. Asclepias, Euphorbia, Lysimachia, 
Peonia. Tournefort and his contemporaries resumed this 
practice, and named plants in memory or in honor of distin- 
guished botanists. Ex. Begonia, Bignonia, Cesalpinia, Fuchsia, 
Gerardia, 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 
adherentibus ; 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 ‘* differentia,” thus used as specific names (the 
nomina specifica of Linnzeus), became extremely cumbrous. It 
was about in the middle of his career that Linnzus suggested 
what he called frivial names (nomina trivialia) for the specific 
name, consisting of a single word; and in the Species Plan- 
tarum, in 1753, he carried this idea into full effect in Botany. 
The step was a simple one, but most important; and Linnzeus 
himself. who generally did not underrate his services to science, 
seems hardly to have appreciated its practical value.* 

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 Grown or Jones; the specific answers 
to the baptismal name, as John or James. Thus, Quercus alba is 
the botanical appellation of the White Oak; Quwereus being that 
of the genus, and ala (white) that of a particular species ; while 
the Red Oak is named Quercus rubra ; the Scarlet Oak, Quercus 
coccinea; the Live Oak, Quercus virens; the Bur Oak, Quercus 
macrocurpa: Magnolia grandiflvra is Large-flowered Magnolia ; 
M. macrophylla, Long-leaved Magnolia, and so on. The name of 
the genus is a substantive, or at least is a word taken as a substan- 
tive. That of a species is mostly an adjective adjunct, always 
following the generic name and in the same gender.?. This com- 
bination of generic and specific name is the name of the plant. 

705. 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,* 


1 Moreover, he may be said to have adopted rather than originated the 
idea; for single-worded specific names were used half a century previous 
by Bachmann, a/ias Rivinus. 

2 It is to be noted that the classical Latin names of trees are all feminine, 
therefore Quercus alba, Pinus rigida, &e. 

8 The name of a Sean. is sometimes written in between the two parts 
of the plant’s name, as Prunus (Padus) Virginiana. This is the name of the 
plant and something more. In addition to the name of the species, that of 
the variety or even subvaricty is sometimes added. 

# Alphonse DeCandolle several years ago estimated the known species of 
Flowering Plants at between 100,000 and 120,000. The larger number may 
perhaps include the higher orders of the Flowerless series. In the present 
state of our knowledge of the lower orders of Cryptogams, no close estimate 
can be well formed of the actual number of species. 


NOMENCLATURE. S47 


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 system 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 Linnzeus, 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 Théorie Elémentaire) by DeCandolle 
‘* who was ruled by the idea of having the law of priority prop- 
erly respected,” was critically considered by Lindley 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.! 

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, ete., Geneva and Paris, 1867. 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., 1868. 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, 
1877 ; also, Bentham in Journal of the Linnean Society, xvii. 189-198, 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, 1879. 

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 PHYTOGRAPHY. 


that may 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. DeCandolle, |. ¢.) 

708. The following are universal rules in scientific nomen- 
clature : — 

1. Names must be in Latin or be Latinized. Those from the 
Greek (which are more and more abundant, owing to the facility 
of this language for compounding) take Latin form and termina- 
tion. Those from modern or other than classical languages 
should at least have a Latin termination.2 Hybrid names, 
namely, those formed by the combination of two languages (at 
least of Latin and Greek), should not be made.? ; 

2. For each plant or group there can be only one valid name, 
and that always the most ancient, if it is tenable. 

3. Consequently, no new name should be given to an old plant 
or group, except for necessity. That a name may be bettered 
is no valid reason for changing it. 

709. Names of Genera are substantive and singular, of one 
word; and the same name cannot be used for two genera of 
plants.* They may be derived from any source whatever, from 


1 Thus, words ending with the Greek os generally change it to us, and with 
ontoum. <A rule not always observed; for while we have Epidendrum and 
Oxydendrum, Linneus himself variously wrote Liriodendrum and Liriodendron, 
Rhododendrum and Rhododendron ; and the Greek form now prevails. 

2 In this as in other cases, some exceptions are well established by 
custom, but they ought not to be extended. The rule as to Latinization is 
restricted as respects orthography by the necessity of preserving modern 
commemorative names in a recognizable form. 

3 But we cannot change numerous old names for this fault, such as con- 
voluloides, fumario‘des, ranunculoides, and scirpoides (though they ought to 
have been convolrulina, ranunculina, and scirpina) ; and modern botanists have 
not scrupled to append the expressive and convenient Greek term -o/des (sig- 
nifying likeness) to generic names not of classical origin. Ex. abutiloides, 
biroides, davallioides, fuchsioides, gentiunoides, lobelioides, tournefortioid:s. In Eng- 
lish, some hybrids will perpetuate themselves, as for instance lerminology, 
centimetre, millimetre, beaurocracy, &e. 

4 Very many, indeed, are adjectives used as substantives, as Arenaria, 
Clararia, Saponaria, Impatiens, Trientalis, and even Gloriosa, Mirabilis, &e. 

Some two-worded generic names anterior to Linnzus, such as Dens Leonis, 
Vitis Idea, Bursa Pastoris, remain for sections and species, but not for gen- 
era. When two words are confluent into one, they are not objectionable, 
as Laurocerasus, Carlemania (commemorating Charles Leman, Carolus Le- 
manus), &c. 


NOMENCLATURE. 3840 


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 combins- 
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 they should be adaptable 
to the Latin tongue. Characteristic names, when possible, are 
among the best; such as Sungucnaria for an herb with red juice, 
Hemutoxylon for the Logwood tree, Lithospermum for a plant 
with stony seeds (or seeming seeds), Myvsurus for a plant with 
gyneecium 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, 
Polygala is wholly destitute of milk, and many species of Con- 
volvulus do not twine. Neat anagrams are not bad, such as 
Brown’s 7el/ima 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 Linnzeus and Jussieu downward. In 
forming such names, the name of the person, cleared of titles 
and accessory particles (thus Cundollea, not D.candollea), takes 
the final -a or-/a and becomes feminine ; and its orthography is 
preserved as far as possible, making only necessary concessions 
to euphony and to the genius of the Latin language.? 


The Linnean 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 phenogamous names in the lower Cryptogamia. 

1 Thus, we may write Lescuria instead of Lesquereurra, although Michauria 
is the form for the genus dedicated to Michaux, however pronounced. The 
genus dedicated to Strangways is written Stranvesia (although Strangwaysia 
might have been tolerable) ; to Andrzeiowsky, Andreoskia ; to Leeuwenhoek, 
Levenhookia (although the elder DeCandolle restored all the vowels), &e. 
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 Lechenaultia 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 misunder- 


350 PHYTOGRAPHY. 


710. The etymology of a new genus should always be given. 
Of the Linnzan restrictions, one holds, viz. that the names of 
genera are not to end in -odds, 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, habit, 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 
Ranunculus, R. bulbosus is named from the bulb-like crown or base 
of the stem; &. acris, trom the acridity of the juice; J. scede- 
ratus (the accursed), in reference to the same property ; 2. repens, 
from the creeping habit of the stems; &. pusilius, from general 
insignificance ; /. aquatilis, from its growing in water; R. ni- 
valis, from living near eternal snow; /. Pennsylvanicus, from 
country or State whence it was first made known to botanists ; 
fh. 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 Nuttaliii, ¢. e. the Ranunculus of 
Nuttall, instead of 2. Nuttallianus, the Nuttallian Ranunculus. 
Yet the latter form is preferred when the species is named in 
honor of some one who did not discover nor treat of it (which 
should seldom be) ; but this distinction is a custom rather than 
a rule, and the form of the commemorative name may be settled 
by euphony or convenience. In any case, the personal name 
should have a capital initial. 

712. Many specific names are substantives, occasionally a 
common substantive, as Stellar/a nemorum (of the groves), Con- 
volvulus sepium (of the hedges), Cassia pumilio (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. 
Ranunculus Flammula, R. Thora, and R. Cymbalaria; 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 Greek 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 Tournefort, Linneus, Jussieu, and their contemporaries, has by pre- 
scription rights which botanists are bound to respect. Wherefore Pyrus 
is the botanical name of the pear-tree, notwithstanding the classical Pirus. 
So Jervis, as a specific name for a smooth plant (and as distinguished from 
lévis, a light or slight one), is fixed by long botanical use, although only 
lévis 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-caruleum; and some of. ancient 
origin (once quasi-generic) are of two words. Ex. Panicum 
Crus galli, Cupsella Bursa pastoris, 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 by 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 Flammula, var. reptans, and FR. agquatilis, var. tricho- 
plyllus. 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 B, 
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 & prefixed. Ex. & Salix capreola. Hybrids of known 
parentage are named by combining the names of the two pa- 
rents, thus: S. purpureo X daphnoides, or K S. purpureo-daph- 
noides, for a cross between S. purpurea and 8. daphnoides, of 
which the first supplied the pollen to fertilize the second. The 
counterpart hybrid is & S. daphnoideo-purpurea. 


1 In respect to the initial of geographical specific names, being adjec- 
tives, such as Americana, Canadensis, Virginiana, Europea, Anglica, 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 usua! 
practice to write such geographical specific names with a small initial. 


b02 PHYTOGRAPHY. 


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.! 

719. A species is not named unless it has assigned to it both 
a generic and a specific 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 /xrsiccate in place of printed descriptions), is held to 
be tantamount to publication.? 

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 ‘* Rununculus pratensis erectus acris,” according to Bauhin, 
in his Pinax, p. 179. Under the new nomenclature, which re- 
duced the specific part of the plant’s name to one word, this 
became Ranunculus acris in Linnzeus, Species Plantarum (ed. 1), 
p- 904; and a brief character gave its distinctions. In later 
works it has been more fully dcseribed, 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 be communicated, in manuscript or otherwise, by the pro- 
pounder to an author who may make them known by publication; but the 
date of the genus or other group is that of actual publication. 

2 This does not cover all the conditions of publication, since it does 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 are not in the category of “ unpub- 
lished names,” which generally ought to be left untouched. 

8 For good examples of bibliogranhy, see such detailed works as De- 


NOMENCLATURE. 30e 


anist, in referring to this or any other plant, might cite any work 
which describes it, or none at all. Ranunculus aeris 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 abbreviated name of 
Linnzeus.! Here we have the name of the plant, and the bibli- 
ography reduced to its initial. To this, further citation and 
other references may 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 Linnzus 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 may have differed from the earlier 
in his views, and have referred the plant to some other genus. 
As instances of the first, Huphorbia 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 naine 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 7. Darlingtonit, in honor of the discoverer and first 
describer. The common Milkweed of Atlantic North America 
was named by Linnweus Asclepias 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. Cornuti, in commemora- 
tion of an ante-Linnzean 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, 
Z., meaning an Anemone resembling a Thalictrum. 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 
done in Michaux’s Flora; and so the accepted name is Thalictrum 


Candolle’s Systema Vegetabilium, and Sereno Watson’s Bibliographical 
Index to North American Botany, in the Smithsonian Miscel!aneous Col- 
lections. 

1 For Abbreviations of Authors’ Names, see 385. 


304 PHYTOGRAPHY. 


anemonotdes, Michx., meaning an Anemone-like Thalictrum, and 
Michaux is the authority for this name. The names which for 
any reason are superseded become Synonyms? (75d.) 

722. A later author may circumscribe a species or a genus 
differently from the originator of the name. To a greater or 
less extent, this must continually happen in the course of time. 
But ** Ricinus communis, Linn.,” stands unmoved by the sub- 
sequent admission of various species (known or unknown to 
Linneeus) and the final reduction of all to one by a thorough 
monographer. So does Silene Gallica, Linn., although S. guin- 
guevulnera, Linn., of the same date, is reduced to it. There is 
no suflicient reason for writing Myosotis, Brown, or Cynoglossum, 
Brown, because this author restricted the limits of these genera ; 
nor to write Gilia, LBenth., because Bentham vastly extended 


1 The synonymy is an essential part of the bibliography or scientific 
history of a genus or species. But synonymous and admitted names ought 
to be kept distinct. 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 the 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 zoologists and some botanists, 
and which has of late years been very fully discussed. 

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, 
Leontice thalictroides, Linn., 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. P/. 312,” and might be further ex- 
tended. In the Flora of Michaux, this plant was treated as distinct from 
Leontice in genus; and some botanists adopted this view, while others of 
equal authority did not. Those who adopt Michaux’s genus name the plant 
Caulophyllum thalictroides, Michz. 

Now 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: Caulophyllum thalictroides, 
Linn. Or else they would avoid direct falsification of the facts by adding 
(sp.), this being explained to mean that the specific part of the name only 
was given by Linneus. Then, as this omits all mention of the original gen- 
eric part of the name, others add this ina parenthesis, and write: “ Caulo- 
phyllum thalictroides (Linn. sub Leontice) Michr.,” or “ Caulophyllum (Michx.) 
thalictroides, Linn. sub Leontice,” or “Caulophyllum (Leontice, Linn.) thalictroides, 
Michr.”’ 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 brevity of the binomial nomenclature. It is all but certain that, 
if the genus Caulophyllum had been published in the lifetime of Linnzus, 
he would not kave 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. sp.,” 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.’ 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 synonymous 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. simpler, Torr. § Gray, and not of 
Michaux, for it is the species P. simpler, Michr. Geum, subgen. Stylipus, 
Torr. & 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, Uriseb ;” 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. 
Sperguaria 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 
Chilopsis 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 Coir arundinacea, then named 
by Roxburgh as Coir larbata, and entered in Sprengel’s Systema with 
Willdenow’s character as Coir Kenigii. All these names were defective 
as referring to a wrong genus. Brown corrected the error by creating the 
new genus Chionachne, and selected Roxburgh’s specific name as the one 
most generally known and the least liable to misinterpretation , and Brown’s 
Chionachne barbata is therefore the first correct name; for which Thwaites 
afterwards substituted Chionachne Kanigii, 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- 
acez, in Jour. Linn. Society, xvii. 197, 198, November, 1878). The following 


3856 PHYTOGRAPHY. 


725. Names of Subgenera or of other sections of genera are 
like those of genera; indeed very many of them, and the most 
fitting, are eld generic names which haye been comprehended 
in the genus by reduction. Unlike genera and higher groups, 
however, sections, when of Greek derivation, may properly take 
the termination in -ovdes,t and the typical section may bear the 
name of the genus with the prefix Lu.” Sections need not be 
named at all, and only those of comparatively high rank should 


is a farther extract from the same protest against the practice ‘of creating 
a new 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 
the 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 specific name a 
general substantive aspect, as if the generic ones were mere adjuncts, — 
a serious encroachment on the beautiful simplicity of the Linnean nomen- 
clature; and it is to be feared that there is a tendency in that direction in 
phenogamic 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 genus. It is 
probably from not perceiving the difference between making and changing 
a name that the practice objected to has been adopted by some of the first 
among recent botanists.” Bentham, I. e. 

1 A genus could not properly have one of its sections called by its own 
name with the addition of -o/des or -opsis, as Asteroides or Asteropsis, for it 
is senseless to declare that an Aster resembles an Aster; but sectional names 
of this composition may be excellent for sections of other genera, as ex- 
pressing analogy or resemblance. Latin generic names used for sectional 
ones properly take the addition of -e//a, or -ina, or -astrum. 

2 The prefix Lu (Greek for much, very, or true), prefixed to a generic 
name of Greek origin, is the proper designation of the typical section of 
that genus, meaning the group which should bear the generic name if such 
genus were divided. The rule against hybrid names should in strictness 
exclude this prefix from Latin names, but it has not always done so. 


NOMENCLATURE. 307 
have substantive names. Designations, however, are conven- 
ient for lower sections; and the name of a leading species 
may be used, in the plural; as Aster, section Amelli, and sect. 
Concinni. Subgenera need not agree in gender with the genus 
they belong to. When written with the name of the plant, 
the subgeneric name is parenthetically inserted between the 
generic and specific appellation. Ex. Pyrus (Malus) coronaria. 

726. Names of Tribes, Orders, &c. The names of all groups 
superior to genera are adjectives plural, and with few excep- 
tions are the names of genera lengthened by some adjective 
termination. Ex. From Rosa, Rosee, Rosacee, Rosales; from 
Myrtus, Wyrtee, Myrtaceae, Myrtales ; from Berberis, Lerberidee ; 
from Tamarix, Zumaricinee ; from Salix, Sulicee, Salicinee. 
The substantive Plante being understood, the groups are Rose- 
ous, or Rosaceous, or Rosal plants, &e. 

727. Tribal Names, and names of whatever grade between gen- 
era and orders, are formed by adding to the root of a generic 
‘name a final -e@. Ex. Rosee, Phasevlee, Antirrhinee, Oxalidee, 
&¢. Some subtribes take the name of the tribe with the prefix 
Eu, as Eiphaseolee tor that subtribe of the tribe Fhaseolez 
which comprises the representative genus Phaseolus. Tribal 
names may take the same prefix, as Huc@salpinee for the tribe 
of the suborder Ceesalpinez which contains the typical genus 
Cvesalpinia. 

728. Ordinal Names are formed in the same way, but with a 
preference for certain terminations which may denote their rank, 
especially that of -acee,— as Rosacea, Myrtacee, Cucurbiiacee, 
meaning Rosaceous, Myrtaceous, and Cucurbitaceous plants. 

729. The names of what we now call natural orders, as 
sketched or adopted by Linnaeus, were mostly descriptive, such 
as Ensate, Spathacee, Coronarie, Pupilionacee, Conifere, Amen- 
tacee, Umbellate ; but a few took their names from genera, as 
Orchidee, Liliacee. Jussieu, with whom the system of natural 
orders properly began, had no suborders, tribes, or any such gra- 
dation of groups to deal with. His one hundred ordinal names 
are some of them of the descriptive kind, as several of the above, 
also Leguminose, Corymbifere, &c. But the greater part are 
simply plurals of generic names, such as Asparag/, Junci, Lilia, 
Muse, Orchides, Lauri, Convoleuli, Erice, Acera, Cacti. To a 
few was given the lengthened termination in -ee@, as Polygonee, 
Solanee, Berberidee, Curyophyillee ; to some, the termination in 
—ace@, as in Cichoracee, Campanulacee, Leubiacee, Ranuncu- 
lacee, Malvaceae, Tiliacee, Cucurbitacee. Subsequent authors 
have necessarily changed all names which were plurals of gen- 


358 PHYTOGRA PHY. 


era; and the strongly prevalent tendency has been to give the 
termination in -acee to all such ordinal names, and to restrict 
this termination to orders. Lindley insisted upon making this . 
an absolute rule even for names not formed from generic appel- 
lations: but this will not be adopted. 

730. In the first place, several large orders which have been 
known from the first by such characteristic names as Crucifere, 
Leguminose (and its suborder Papilionacee), Guttifere, Umbelli- 
fere, Composite, Labiate, Cupulifere, and Conifere, also Palmee 
and Graminee, Filices, and even Aroidee and Ficoidee, will retain 
these appellations ; but no new ones of the kind will be made. 

731. Also, names formed from genera which do not well take 
the termination in -ace@ may be allowed as orders to retain their 
natural form in -cnee, -idee, -ariew, and the like. Ex. Tamaris- 
cinee, Salicinee, Scrophularinee, Berberidee, Lentibuluriee. We 
may prefer for the sake of uniformity to write Salicacee, Berberi- 
dacee, Lentibulariacee, and Scrophulariacee (as we should write 
Violacee), but this form cannot be insisted on. On the other 
hand, a termination in -acee 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 
Vernoniacee, to avoid Vernoniee, which ends with four vowels. 
Spireea and Staphylea are the types of tribes, for which the names, 
if they followed the rule, would be Sprreee and Staphyleee, 
ending one in five the other in four consecutive vowels. Some 
avoid this by writing Staphyleacee and Spireacee. Others write 
Stuphylee, but this is only the plural of the generic name. 

752. A few orders or other groups took their names long 
ago from superseded generic names. Ex. Caryophyllacee or 
Caryophyllee, Onagracee or Onagrariee, and Lentibulariee. 

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. Ranales, 
Parietales, Malvales, Rosales, Passiflorales, &c., most of them 
founded on the names of representative genera and orders. 
Euphony requires some to take other terminations. Ex. Poly- 
galine, Caryophylline. 

734. Names of Classes and other great divisions are plurals, 
either adjective or adjective nouns, expressive of the leading 
character. Ex. Polypetale, Gamopetale, Apetale ; Angiosperme 
and Gymnosperme ; Dicotyledones and Monocotyledones. The 
names of the two great series or sub-kingdoms, following the 
analogy of the Linnzean classes, end in -ia, and are Phenogamia, 
or Phanerogamia, and Cryptogamia. 


GLOSSOLOGY. 359 


Section II. Gtrossotocy or TEermMiInoioey.? 


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 Linnzeus, and then to DeCandolle. 
The Théorie Elémentaire 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 
ful in phenogamous 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 by 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, howev er 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 absolutely 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 ina 
Ranunculus, a follicle in Aquilegia, a berry in Hydrastis and 
Actzea; 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 répua, of the same meaning. 


260 PHYTOGRAPHY. 


stantive instead of merely adjective names. But the former 
should not be unnecessarily multiplied. 

738. The classical language of scientific botany 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. Greatly to its advantage, 
English botanical terminology has adopted and incorporated 
terms from the Latin and Greek, with slight changes, not obscur- 
ing the identity, thus securing all their precision, and rendering 
the simple botanical Latin of descriptions of easy acquisition to 
the 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), are 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 DeCandolle 
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. 

741. 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- 
lanceolate, for an outline between the two. Also a term may be 
qualified by the prefix sb, in the sense of somewhat, as in sud- 
rotundus, subcordatus (somewhat round or slightly heart-shaped), 
or diminutives (such as ¢ntegriusculus), or superlatives (cnteger- 
rimus) or Other strengthened forms (such as perangustus) may 
be employed. Among terms of more than one form of meaning 
are such as calycinus. which may mean, according to the context, 
pertaining to the calyx, or of the appearance of calyx; cymosus 
may mean in cymes, or bearing cymes, or in the manner of a 
cyme; and paleaceus may mean provided or beset with chaff, or 
resembling chaff in texture. Often the form of the word should 
distinguish the sense ; as fol/atus, furnished with leaves, foliosus, 
with abundance of leaves, while foliaceus may mean either bear- 
ing leaves, or properly of leaf-like texture or appearance. 


DESCRIPTION. 361 


742. Absence of an organ or quality may be expressed by 
means of a prefix with privative signification, as trdehiscent, not 
dehiscent, exannulute, destitute of a ring, apetalous, without 
petals. But the Greek privative « should not be prefixed to 
Latin words, nor the Latin sud to terms taken from the Greek. 

743. When the Latin preposition 04 is prefixed to an adjective 
term, it means obyersely ; 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. DeEscriIprion. 


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- 
nus, 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 solely. 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 differenti@, ov 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 may 
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 The former would answer to what have been termed d/fferential char- 
acters, the latter to essential characters. Linnzeus 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 Linneus and his predecessors; what we call the specific, 
Linneeus called the trivéal name. (703.) 

747. Subordination of characters and the avoidance of vain 
repetitions require that as far as possible — regard being had to 
the form of the work —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 
the generic character that of the superior groups ; and so of the 
sections and subdivisions of all grades down to the species. 
Equally from the specific character should be excluded every 
thing which belongs 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 varicties 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. What 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 writers 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. Rubiacez, for example, are known by their opposite 
entire and simple leaves and intervening stipules, along with a 
few floral characters; Sarraceniaceze, by tubular or pitcher-like 
leaves, along with a certain combination of a few other charac- 


ficially together, as they might be in an artificial key, and as very unlike 
genera often were in his sexual system; by the second meaning the distine- 
tions, the fewer the better, which will separate a group from its nearest 
relatives ; by the third, all real marks of difference, 7. e. all afforded by the 
organs of fructification, which only were taken into account for genera, &c. 
Upon the construction of this natural character Linneus prided himself, 
and justly. These are the characters in his Genera Plantarum. 


CHARACTERS. 503 


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 may be 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, gyncecium and its parts, 
ovules; then the fruit, seed, albumen, if any, embryo and its 
parts. But descriptions of this sort in most works and in ordi- 
nary cases are partial and subsidiary, comprising only 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 by 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. 

751. Linnzus required that neither the essential character of 
a genus, nor a specific character (his nomen specifieum), 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 
brevity, or such as the case admits, is secured by proper group- 
ing under a subordination of sections, subsections, and other 
subdivisions.” 


1 Ex. “Nepera Cataria: erecta, elata, cano-pubescens ; foliis petio- 
Jatis,’” ete. In English, these adjectives without any substantive expressed 
will be seen to belong, as here, to “ plant” or “ herb ” understood. 

2 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 


504 PHYTOGRAPHY. 


752. Punctuation. In proper descriptions, and in characters 
of genera and of 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. Excepting the latter part, the adjective terms 
are separated by 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. Subgeneric 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 characiers are replaced by a characteristic 
synopsis at the head of each genus; and a terse description under each 
species completes the account. Moreover, Bentham, in recent works, such 
as his revision of the Genus Cassia, also that of the Mimosez, which have 
Latin characters, writes these in the nominative case and cach member in a 
separate sentence, in the descriptive form, abandoning the long-used abla- 
tive form. 

1 Linneus 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 connna. Thus, 

“CHEeNOPoDIUM ALBUM foliis rhomboideo-triangularibus erosis postice 
integris : summis oblongis, racemis erectis.” Spec. Pl. ed. 2, 519. 

Here, while the two main members of the sentence are separated by a 
comma, a subsidiary portion of the first member, relating to the uppermost 
leaves, is separated by a colon. Linnzus employed the colon in the same 
way in generic characters. This anomalous usage is now abandoned. But 
most authors have followed the Linnean 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, 

““Ranuncutus acris (Linn. Spec. 779) foliis pubescentibus subglabrisve 
palmato-partitis, lobis inciso-dentatis acutis, summis linearibus, caule erecto 
plurifloro subpubescente, pedunculis teretibus, calyce subvilloso, carpellis 
mucrone suberecto 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 characterize 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 fer the 
secondary ones,— a punctuation now not uncommon, and which is adopted 
in the recent first volume of the Monographie Phanerogamarum of the De- 
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 abiative phrase (as, 
“Smilax laurifolia ; limbis foliorum oblongis vel ovato-oblongis, coriaceis, 
3-5-nerviis, subtus pallidioribus,” etc.), where they are generally deemed 


PUNCTUATION, SYNONYMY. 369 


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 Linnei. Still, since when the suthor’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, Gray, Ann. N. Y. Lye. 3. 221. Torr. & 
Gray, Fl. 1. 113,”? &e. 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 partly under the admitted name, 
and partly in the 

Zoo. 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.  Nectr/s 
of Schreber is a synonym of Cabomba of Aublet, the latter hay- 
ing priority.  Thalictrum anemonoides of Michaux has for syno- 
nyms Anemone thalictroides of Linneus and of many subsequent 
authors who followed him in referring this ambiguous plant to 
Anemone (721) ; and also Syndesmon thalictroides of Hottmannsegg 


5) 


and Anemonella thalictroides of Spach, who proposed to consider 


superfluous. The preferable punctuation of the character above-quoted 
from the Prodromus would be 

Ranuncucus acris (Linn.): foliis pubescentibus subglabrisve palmato- 
partitis, lobis inciso-dentatis acutis, summis linearibus; caule erecto pluri- 
fioro 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 North 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. 


306 PHYTOGRAPHY. 


it an intermediate genus between Thalictrum and Anemone. In 
systematic works, the specific character immediately follows the 
name, and generally forms a part of the same sentence ; and is 
followed first by citations of authors who have adopted the name, 
and then by the synonymy, or as much of it as the plan of the 
work calls for. The synonymous 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 date. Also, where the synonymy is not elaborately 
displayed, the various synonyms of the same generic name are 
usually placed in consecutive order. 

756. Ieonography. ‘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 figures 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. 

757. Habitat and Station are recorded in a sentence or para- 
eraph following the name, character, and synonymy of a species. 
The habitation 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 affects, whether in water, in marshes, on shores, 
on hills or mountains, in forests, on open plains, &e. 

758. Discoverer, &e. To the habitat and station of newly 
discovered, rare, or local plants should be appended the name 
of the discoverer or the collectors by whom the species has be- 
come known to science, at least when the plant is first published. 
Date of discovery should also then be indicated. 

759. Time of Blossoming 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, flowering-time is intended. In a flora, this may 
sometimes be indicated under the genus for all the species. 
In the flora of an extensive region, and in respect to species of 
considerable range in latitude or longitude, the time of flowering 
differs so widely at the extremes of the geographical range that 
it cannot well be specified except in general terms, as spring, 
summer, autumn, &e. 

760. Etymology 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. Aecentuation 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 syllable upon which the principal accent 
falls. This in, words of two syllables is always the first; in 
words of three or more syllables, 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 Eyica, or short as in Arbaus and Gludidlus. Or 
else the accent may be marked by a proper sign, as Erica, Arbutus, 
Gladéolus. An endeavor has been made to represent the longer 
sound of the vowel by the grave accent-mark, as Hrica, and the 
short 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 Mooker), or the 
first two letters following the vowel when both are consonants 
(as Linn. for Linnzeus, Juss. for Jussieu, ich. for Richard). 
Sometimes more of the name must be given, in order to distin- 
guish those beginning with the same syllable. So we write Michr. 
to prevent confusion of the name Michaux with that of Micheli, 
which, being the earlier, claims the abbreviation M/ch., 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 Juss. fil., Hook. fil. 
or Hook. f., &¢. Certain, but very few, well-known and eminent 


268 PHYTOGRAPHY. 


names are abbreviated to a sign; as L. for Linnzeus, DC. for 
DeCandolle, HBik. for Humboldt, Bonpland, and Kunth, the 
latter too long after ordinary abbreviation.’ Care should be 
taken to afiix the period by which abbreviations may be dis- 
tinguished from full names, such as Don, Ker, blytt. 

764. Abbreviations of titles of works follow the same rules as 
those of names, or at least are in no wise peculiar in botany. 

765. Abbreviations of the names of organs follow the same 
rule: Cal. for calyx, Cor. for corolla, Stam. for stamen or 
stamina, st. for pistillum or pistil, “r. for fructus or fruit, 
Per. for pericarpium or pericarp, Sem. for semen or seed, are the 
most common. ab. for habitat or geographical station, Herd. 
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 least 
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, v. s. s., when it is a spontaneous specimen 
that has been examined in a dried state, and v. s. e., 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 are 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. See p. 391. 

767. The marks used 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, 
S$ * + + =. The first one, for sections of the highest order, 
takes numerals after the sign. Ex. § 1, and so on. When 


' As Alph. DeCandolle remarks, the proper abbreviation of the name he 
bears is Cand. But the form DC. was very early adopted by the first of 
the illustrious name, and has been continued for almost three quarters of a 
century. Alphonse DeCandolle would prefer to write it D.C., but has not 
adopted that mode, nor should we; for DC. and HBK. are convenient ab- 
breviations reduced to signs. But such forms should not be increased. For 
ordinary names they would be unintelligible. 

Names which are not too long, and of which an abbreviation by the ordi- 
nary rule is insufiicient, 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.’ Ex. Phacelia, Juss., § 1. Euphacelia, 
?.e. the true or typical Phacelia; § 2. Cosmanthus, Gray, Ke. 
Sections next in rank to these are marked with asterisks, * for 
the first, * * forthesecond, * * * forthe third one of the same 
rank. Divisions of these have the + prefixed ; and so on in the 
sume way. Still farther subdivisions may be marked by the 
small letters of the alphabet consecutively. a, d,c. When capital 
letters are used for division marks, it is mostly for those of a 
high grade. 

768. Floras, Monographs, &e. A systematic work describing 
in proper order the plants of a country or district is generally 
alled a Flora, A Flora of a small district takes the diminutive 
name of a Mlorula. <A universal work of the kind when it ex- 
tends to the species is a System, Systema Vegetabilium or Systema 
Regni Vegetabilis. The latest completed Systema Vegetabilium 
is that of Sprengel (1825-1828), in five octavo volumes, on 
a very condensed pian. 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 
by the same author. The principal work of this kind is the Pro- 
dromus Syst. Nat. Regni Vegetabilis, commenced by DeCandolle 
in the year 1824, continued by his son Alphonse DeCandolle 
(aided by various botanists) to its close in 1873, down to vol. 
XVii., or essentially twenty very compact octavo volumes, these 
carrying the work only through the great class of Dicotyledones. 
But the publication of the monocotyledonous orders has com- 
menced in a series of Monographs (Monographie 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 Jussieu, 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, &e., 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, &¢. 


770. Implements. Those necessary for the examination of 
phenogamous 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. Forasingle 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 (doublets 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 scalpél 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 system. 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 anthotaxy or 
inflorescence is to be examined and referred to its proper type. 
In the flower, 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 estivation 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 dehiscence, &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. atk 


determined. Two sections of the flower should be made: one of 
them vertical and directly through the centre, in the manner of 
Fig. 336-341, —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 zestivation. (Fig. 351, 886, 398, 
&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 embryo 
may almost always 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 
they may be made to express the whole floral structure, even to 
the coalescence and adnation.' 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 essentially 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 may be. 

776. Herborizing.? The collector’s outfit will essentially con- 
sist of a Vaseulum or botanical box, a Portfolio, a Trowel, a pocket 
Lens, and a small but stoutly covered Note-book. Some use a 
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 Eichler’s Bliithendiagramme (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. Hoysrapt, 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. 


at2 PHYTOGRAPHY. 


ciass. 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 be kept meanwhile under some pressure. 

777. The Vaseulum is very useful for holding plants that are 
to be examined fresh, and for thick roots, large fruits, &ce. It 
is made of tin, and should be of oyal-cylindrical shape, about 17 
inches long and 4 by 6 inches wide, and provided with a light 
strap to throw over the shoulder. The lid opens nearly the 
whole length of one of the flat sides (15 by 44 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 
may 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 
very 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 11} inches may be better, as 
there would then be little danger of making specimens of too 
great length for the herbarium. (784.) Or the 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 exceilent for this purpose, being sufficiently 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 are 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 drier to another in the drying process. 

780. For digging up roots, bulbs, &c., a small and sharp 
pointed triangular Zrowel 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.’ 

781. A Note-book should be carried upon every 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 
memory. 

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 Cruciferze, the Umbellifere, 
and the Cyperaceze. With many plants the radical-leaves, with 
others the character of the subterranean stem, whether a rootstock, 
tuber, corm, 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 vite 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 dicecious shrubs or 
trees, like the Willows, each species should be represented 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. 

784. 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. 

785. Carices should be always collected when the fruit is full- 
grown, but not so ripe as to fall away. So also should other 
Cyperacee ; 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 
difficulty is promptly remedied by crushing 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. 3875 


786. Some aquatic plants (Algz 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, cither 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 directly 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 
Lycopodium 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 locality loses much of its value and interest. 

788. Drying Specimens. ‘The chief requisite for good herba- 
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, called 
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 may 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 


310 PHYTOGRAPHY. 


evenly distributed. Hence it is best to have these sections not 

over five or six inches in thickness, nor should the pile itself . 
be carried too high, never exceeding two feet. Painted binders’ 

boards may 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. This is 

said 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 haye been contrived. 
The Serew-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 Lever-presses, but 
they are usually unwieldy. Fortunately, one of the best forms 
of the drying-press, as well as the simplest and cheapest, is 
merely a board with weights placed on the top of the pile of 
specimens. Here the pressure is continuous, constantly follow- 
ing the shrinkage of the plants. The weight ona pile should 
vary from 25 to 100 pounds, according to the nature of the 
specimens and the quantity in the press. On an average, 60 
pounds is sufficient for most plants. If much greater pressure 
is used, there is danger of crushing the more delicate parts of the 
specimen, and thereby impairing its scientific value. For weights, 
bars or masses of iron may be used, boxes filled with sand, 
stones, and the like. 

791. Specimens brought home in the botanical box must be 
placed in such thin specimen-sheets as are used in portfolio. 
In putting plants in specimen-sheets, whether in portfolio or 
press, it is well to take some pains to spread out the specimens 
neatly ; for a little care now may save much later labor. How- 
ever, with most species, any carelessness in this respect can 
be remedied at the first change of driers. But there are some 
plants, previously referred to, so peculiarly sensitive that what- 
ever adjustment they receive must be given at the time they are 
first placed in their sheets. 

792. Aithough plants can, if necessary, be kept fresh for several 
days in box or portfolio, on returning from a collecting trip they 
should be transferred to the home press as early as possible. In 
the transference, particular care should be taken to straighten 
out and remove all folds and crumpling of the leaves, petals, 
fronds, &c., and to arrange the specimen as naturally as possi- 
ble, so as to show the proper habit. Both sides of the flowers 
and leaves should be exhibited. Plants that were put directly 
into press should receive this special attention at the first 
change of driers. which on this account should be made within 


» Ti 


HERBORIZATION. 377 
several hours afterwards. The stubbornness and elasticity, so 
troublesome in specimens when first put in, will then have 
mostly disappeared, and the whole specimen will be found suffi- 
ciently flaccid to have every part stay as arranged. If this first 
change is deferred longer than ten or twelve hours, the speci- 
mens of many species become too dry for making the alterations 
required. At this time small pieces of bibulous paper may be 
placed between leaves, or other portions of the plant which over- 
lap, to prevent moulding or discoloration, and to hasten drying. 
It is well to change these fragments of paper with the driers for the 
first day or two: afterwards they may remain with no detriment. 

793. To have the specimens retain their natural color and 
general appearance, they should be dried as rapidly as possible ; 
and this result is best secured by frequent changing of the driers. 
These should be changed at least once a day for the first four or 
five days, and afterwards every other day, until the specimens 
are thoroughly dried. But a marked improvement in the speci- 
mens will result from more frequent changes during the first 
day or two. The first day with Grasses, Sedges, and their allies, 
and the first two days with most other plants, are of more impor- 
tance than all the subsequent time. As an experienced collector 
declares: ‘‘ Two or three changes of the driers during the first 
twenty-four hours will accomplish more than a dozen changes 
after the lapse of several days. The most perfect preservation 
of the beautiful colors of some Orchids has been effected by 
heating the driers and changing them every two hours during 
the first day.” 

794. Heated driers are very efficient; and the best mode of 
heating is to expose them to the sunshine, and bringing them in 
hot to make the change at once, or as soon as possible. 

795. The number of driers interposed between the specimen- 
sheets should depend upon the plants and the frequency of the 
changes: two will suffice when the driers are changed very often ; 
but more must be employed when the plants are thick and succu- 
lent. Uniform pressure may be secured with large and coarse 
plants by placing strips of pasteboard or pieces of cotton-bat- 
ting about the sides of the package. Ringlets of cotton may be 
placed about some of the larger flower-heads of the Composite, &e. 

796. The time required to dry specimens varies with different 
species and with the season: it depends also on the frequency 
of the changes and the temperature of the driers. By changing 
daily, the time is usually from four or five days to a week. But, 
with two changes a day for the first day or two and with heated 
driers, the process may be completed in half the usual time, and 


3878 PHYTOGRAPHY. 


the specimens will be in much finer condition. An experienced 
collector has no difficulty in ascertaining whether a plant is com- 
pletely cured or not, while to a novice it is often a matter of 
uncertainty. A thoroughly dried plant can be usually told by 
its peculiar hay-like rattle when disturbed; also by placing the 
plant against the cheek. If there is a sensation of coolness, the 
plant is still moist. 

797. If the thick leaves of fleshy plants are immersed for 
a few moments in hot water, the period of desiccation will be 
greatly hastened; but they frequently turn dark as a conse- 
quence of the immersion. The drying of such plants, and 
particularly of the Monocotyledons, may be advantageously ex- 
pedited by placing them between several driers and ironing them 
with hot irons. Small plants may be very neatly dried in old 
books. Very beautiful specimens may be made by placing the 
plant in a tall and narrow vessel, and pouring over it a sufficient 
quantity of clean and dry sand. When the moisture is absorbed, 
it may be flattened in a press. 

798. In shifting the driers of a collection, place the package 
to be changed at the left hand on the table or counter, the new 
pile in front with its length parallel to the person, — a position 
the most favorable for giving any needed attention in arranging 
specimens, — while fresh driers may be placed at the right hand, 
or beyond the pile in front. Thus arranged, the sheets of speci- 
mens can be rapidly shifted into their fresh driers. 

799. The moist driers may be spread out in the sunshine to 
dry, or strung on a line in a warm room, or in the open air, if 
not too windy. Very moist driers may be thoroughly dried 
within an hour, if spread in the hot sunshine. In inclement 
weather, they must be dried by the fire. 

800. To recapitulate the most important points in good speci- 
men-making: Use specimen-sheets to hold the plants undis- 
turbed during the whole process of drying: use plenty of the 
most bibulous driers, sun-dried and heated when practicable: do 
not make the piles too large: make the first shift of driers within 
a few hours, at that time making all needed adjustment of the 
flaccid specimens: change the driers twice a day for the first 
day or two. 

801. For collecting and preserving specimens on a journey, 
or when moving from place to place, some modification of the 
stationary press is requisite. The Z’ravelling-press nmust be porta- 
ble: accordingly the pressure is applied by strong leather straps 
with buckles. There should be three straps, one girding the 
package around each end, and one lengthwise. The top and 


HERBORIZATION. 379 


bottom, if of thin boards, must be cleated, or compounded of 
wood with the grain in opposite directions, or very stout binder’s 
board or trunk-board may be advantageously used. This should 
be covered with coarse cotton or linen cloth, glued fast and well 
painted. While stationary, the pressure may be given by means 
of weights when more convenient. 

802. The Wire press, now much in use, is a press of this porta- 
‘ble kind, in which the boards are replaced by sheets of wire net- 
ting, with wide meshes, and surrounded and strengthened by a 
strong but light iron border. Straps with buckles are used to 
hold the parts’ and contents together and to apply the pressure, 
as in the ordinary trayelling-press. Besides its portability, the 
advantages of such wire presses are that, in a small way, they 
may serve both as portfolio for collecting and as press for dry- 
ing; also that, as the drying takes place mainly by evaporation 
instead of absorption, much less paper is required, and the trouble 
of changing the driers is saved.' In fair weather, the press filled 
with plants may be hung in the wind or sunshine, in foul weather 
near a fire. The disadvantage is that specimens dried in this 
way are apt to be brittle. To use this system advantageously, 
the botanist should have at least two such presses in operation, 
one for collecting, while the other is in use for, drying. 

803. Poisoning. Dried specimens are liable to the depreda- 
tions of certain insects, especially of their larvee. The principal 
pest is a smail brown beetle, Anobium paniceum, L.; the perfect 
insect does considerable damage, the larva vastly more. Plants 
with milky juice, such as Asclepiadezx, Apocynace, and Eu- 
phorbiacez, those containing bitter principles, such as Gentians 
and Willows, and generally such plants or such organs as con- 
tain much protoplasm or azotized matter, are most subject to 
attack. Ranunculacee, Umbelliferze, and Composite are seldom 
spared ; while Labiatez mostly escape, probably on account of the 
volatile oil which they contain. Even Ferns are liable to have the 
parts of fructification eaten away. To a certain extent, the im- 
pregnation of the herbarium-cases with camphor, naphthaline, 
or strong-scented oils, may exclude the vermin. But safety is 
secured only by poisoning. 

804. The proper poison is corrosive sublimate, dissolved in 
strong (95 percent ) alcohol. Drop into the alcohol as much cor- 


1 Prof. A. Wood seems to have been the first to call the attention of 
American botanists to this system, which he has earnestly advocated. 

An improved form of this wire press, well adapted both for collecting 
and pressing in moderate quantity, is made and sold, at a small price, by 
Paul Ressler, optician, at New Haven, Connecticut. 


380 PHYTOGRAPHY. 


rosive sublimate as it will take up, then add a trifle more of alcohol, 
so as to keep the solution just below the point of saturation. 
The stronger the solution the better, except that, at full satura- 
tion and where copiously used, an efflorescence may sometimes be 
left 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 efficient. The moistened specimens should be placed 
between driers and in shallow piles until the alcohol evaporates. 

805. Thoroughly poison all specimens before admitting them 
to the herbarium. It 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 properly dried specimen, duly cared for, should be as lasting 
as the paper which holds it. 

807. The Herbarium, called by the earlier botanists Hortus 
Siccus, 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. 

808. The form of the herbarium as to the size of its sheets is 
considerably variable. That of Linnzeus is of the size of foolscap 
paper: this would now be universally regarded as much too 
small. The principal British herbaria adopt the size of 163 by 
10} inches, 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, 
16} inches in length by 11? 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 164 by 11} 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 ail authenticating tickets 


382 PHYTOGRAPHY. 


or notes, should be attached to the sheet near to the specimen 
they belong to. In view of this, printed and written tickets 
should be of small size.! A ticket which 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. Itis not desirable to parcel out 
the space on a ticket with separate lines and headings for habitat, date of 
eollection, time of flowering or fruiting, name of collector, and the like. 
These 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 roe Herparictm or” 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. 


Sh qprke gia caruka, Tarr. 


tae ae P ar? , le Gee aoe Joe f-¢. aad, 
af J. Jr YAS rag, Aly, 1877. 


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, &e. 


ALPINE FLORA OF THE ROCKY MOUNTAINS. 
Coll. C.C. PARRY. 1872. 


No. IVF 


CoLorabDo, trae 6 Cee 
Geir ove t alt. 2 hip. 


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 ali the species of the same genus, when 
not too numerous, or of a particular section of it, or any conyven- 
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 164 by 113 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 1} or 1? 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: ator 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 away. 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 PHYTOGRA PHY. 


of compartments. Such cabinets can be increased in number as 
required, are portable, and can be disposed in any order, side by 
side or one surmounting another, as may be most convenient. 
The doors should be so constructed as to open and shut readily, 
but to close tightly, so as to exclude dust and insects.? 


1 An excellent plan for small and inexpensive herbarium cabinets, of a 
portable character, is proposed and illustrated by Dr. Parry, in the American 
Naturalist, viii. 471. Each small case is in fact a plain wooden box, wide 
enough to hold two tiers of pigeon-hole compartments, and of any desirable 
height (three compartments high in Dr. Parry’s plan, but double the number 
might be better): the entire front consists of a pair of doors meeting in the 
centre, there fastened by a flush spring catch; the doors bevelled on the inside, 
with a corresponding bevel on the case, to which they are attached by out- 
side hinges, so 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 stand close side 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. 


i. OF NAMES OF BOTANISTS AND BOTANICAL AUTHORS. 


Ach. 
Adains. 


Afz. 


All. 
Amm. 
Anders. 
Andr. 
Andrz. 
Aresch. 
Arn. 
Arrh, 
Asch. 
Aubl. 


Bab. 
Bail. 
Balb. 
Baldw. 
Balf. 
Barn. 
Barr. 
Bart. 

W Bart. 
Bartr. 
Bartr. f. 
Bauh. 
Beauv. 
Benj. 
Benn. 

A. Benn. 
Benth. 
Berg. 
Berk. 


Acharius. 
Adanson. 
Afzelius. 
Agardh. 

C. A. Agardh. 


J. G. Agardh, son. 


Aiton. 
Allioni. 
Amman. 


Andersson of Stockholm. 


Andrews. 
Andrzejowski. 
Areschoug. 
Arnott. 
Arrhenius. 
Ascherson. 
Aublet. 


Babington. 

Baillon. 

Balbis. 

Baldwin. 

Balfour. 

Barnéoud. 

Barrelier. 

Benj. Smith Barton. 


W™- P.C. Barton, nephew. 


John Bartram. 

Wm. Bartram. 
Bauhin. 

Palisot de Beauvois. 
Benjamin. 

J J. Bennett. 

A. W. Bennett. 
Bentham. 

Bergius. 

M. J. Berkeley, 


Berkh. = Berkhey. 


Berken. Berkenhout. 

Berland.  Berlandier. 

Bernh. Bernhardi. 

Bert. Bertero. 

Bertol. Bertoloni. 

Bess. Besser. 

Bieb. Marschall von Bieberstein. 

Bigel. Jacob Bigelow. 

Bisch. Bischoff. 

Behm. Boehmer. 

Boerh. Boerhaave. 

Boiss. Boissier. 

Boland. Bolander. 

Bong. Bongard. 

Bonpl. Bonpland. 

Bork. Borkhausen. 

Borsz. Borszcow. 

Brack. Wm. D. Brackenridge. 

Brebis. Brebisson. 

Bref. Brefeld. 

Brew. &§ | W. H. Brewer & Sereno 
Wats. Watson. 

Brid. Bridel. 

Brong. Brongniart. 

Brot. Brotero. 

Brouss. Broussonet. 

Br., R. Br. Robert Brown. 

PP) Br: Patrick Browne. 

Brunf. Brunfels. 

Buckl. Buckley. 

Bull. Bulliard. 

Burm. Burman. 

Buzb. Buxbaum. 

Cam. Camerarius. 

Camb. 

Ganken: t Cambessedes. 


386 ABBREVIATIONS. 


Campd. = Campdera. Eat. = Amos Eaton. 
Cand. DeCandolle, usually DC. D.C. Eat. D.C. Eaton, grandson. 
Casp. Caspary. Edgew. Edgeworth. 
Cass. Cassini. Edw. Edwards. 
Catesb. Catesby. Ehren. Ehrenberg. 
Cav. Cavanilles. Ehirh. Ehbrhart. 
Cerv. Cervantes. Eichl. Kichler. 
Cham Chamisso. Hiseng. Kisengrein. 
Chapm. A. W. Chapman. Ell. Elliott. 
Chav. Chavannes. Endl. Endlicher. 
Chois. Choisy. Engelm. Engelmann. 
Clayt. Clayton. Engl. Engler. 
Clus. Clusius. Eschs. Eschscholtz. 
Collad. Colladon. Eschw. Eschweiler. 
Colm. -Colmeiro. Ettingsh. Ettingshausen. 
Comm. Commelin. 
Corn. Cornuti. Fendl. Fendler. 
Coss. Cosson. Feuil. Feuillée. 
Cunn. Cunningham, A. or J. Fingerh. ¥ingerhuth. 
Curt. Wim. Curtis. Fisch. Fischer. 
M.A.Curt. M. A. Curtis. Forsk. Forskal. 
Forst. Forster. 
Dalech. Dalechamps. Fourn. Tournier. 
Dalib. Dalibard. Frresen. Fresenius. 
Darl. Darlington. Freye. Freycinet. 
DC. Freel. Freelich. 


{ A. P. DeCandolle. 


A.DC. Alphonse DeCandolle,son. | Goartn. J. Gertner. 
Cas. DC. Casimir DeCandolle, the | Gertn. fC. 'T. Geertner. 


Decne. Decaisne. [grandson. | Gardn. Gardner. 
Deless. Delessert. Garid. Garidel. 
Dennst. Dennstedt. Gasp. Gasparrini. 
Desc. Descourtilz. Gaud. Gaudin. 
Desf. Desfontaines. Gaudich. Gaudichaud. 
Desj. Desjardins. Germ. Germain. 
Desmar. Desmazieres. Gesn. Gesner. 
Desmoul. Desmoulins. Gilib. Gilibert. 
Desv. Desvaux. Ging. Gingins de Lassaraz. 
Dicks. Dickson. Gis. Giseke. 
Diesb. Diesbach. Gled. Gleditsch. 
Dieter. Dieterich. Gleich. Gleichen. 
Dietr. Dietrich. Glox. Gloxin. 

Dill. Dillenius. Gmel. J. G. Gmelin. 
Dillw. Dillwyn. C. Gmel. C.C. Gmelin of Baden. 
Dod. Dodonzus (Dodoens). S. Gmel. S. G. Gmelin. 
D’ Orb. D’Orbigny. Godr. Godron. 
Dorst. Dorstenius. Gepp. Geeppert. 
Dougl. Douglas. Good. Goodenough. 
Dre}. Drejer. Gren. Grenier. 
Dryand. Dryander. Grev. Greville. 
Dufr. Dufresne. Griseb. Grisebach. 
Duham. Duhamel du Monceau. Gren. Greenland. 
Dumort. | Dumortier. Gron. N 


Gronovius. 
Dun. Dunal. Gronov. $ 


ABBREYIATIONS. 
Guett. == Guettard. Jacq. = 
Guib. Guibord. Jacq. f: 
Guillem. |Guillemin. J. St. Hil. 
Guimp. Guimpel. Jord. 
Gunn. Gunnerus. Jungh. 
Guss. Gussone. Juss. 
Adr. Juss. 
Hagenb. Hagenbach. 
Fall. Haller. Kemp. 
Ham. Hamilton. iarct: 
Llanb. Hanbury. Kaulf 
Haust. Hanstein. sents 
Havtin. Hartmann. oe ; 
irschl. 
Hartw. Hartweg. Kit 
Harv. Harvey. K es 
alr. 
Hass. Hassall. Korth 
Flassk. Hasskarl. jon ; 
ostel. 
Hausm. Hausmann. Kremp 
Haw. Haworth. Ve 4 
. romo. 
Hebens. Hebenstreit. fee 
Hedw. Hedwig. ; 
Hegelm. Hegeliaier. 
Hegetsch. egetschweiler. L. 
Heist. Heister. Labill. 
Fleldr. Heldreich. Lest. 
Helw. Helwing. Lag. 
Hemsl. Hemsley. Lall. 
Flenck. Henckel. Lam. 
FHlenfr. Henfrey. Lamb. 
FHensl. Henslow. Lamour. 
Herb. Herbert. Langsd. 
Herm. Hermann. La Peyr. 
Hild. Hildebrand. La Pyl. 
TTochst. Hochstetter. Ledeb. 
Ho fjin. G. F. Hotfmann. Lehin. 
H1. Hoff. Hermann Hoffmann Lem. 
Hoffimanns. Hoffmannsegg. Lesq. 
FHlofin. Hofmeister Less. 
Hohen. Hohenacker. Lestib. 
Holmsk. | Holmskiold. Lév. 
Flomb. Hombron. LD Her. 
Fook. Wm. J. Hooker. LT Herm. 
Hool:. f. J. DY. Hooker, son. Liebm. 
Hopk. Hopkirk. Lightf: 
Hornem. Hornemann. Lili). 
Flornsch. Wornschuch. Lindb. 
Florsf. Horsfield. Lindbl. 
Foust. Houston. Lindenb. 
Houtt. Houttuyn. Lindh. 
Huds. Hudson. Lindl. 
Hueb. Huebener. Linn. 
Hunb. Humboldt. Linn. f. 
HBK. 3 Humboldt, Bonpland, and | Lodd. 


1 Kunth. 


Leff. 


337 


N. J. Jacquin. 

J. F. Jacquin, son. 
Jaume St. Hilaire. 
Jordan. 

Junghuhn. 

A. L. Jussieu. 
Adrien Jussieu, son. 


Kempfer. 
Karsten. 
Kaulfuss. 
Kindberg. 
Kirschleger. 
Kitaibel. 
Keireuter. 
Korthals 
Kosteletzky. 
Krempelhuber. 
Krombholz. 
Kuetzing. 


Linneus. 

La Billardiere 
Lestadius. 
Lagasca. 
Lallement. | Marck). 
Lamarck (Monnet de La 
Lambert. 
Lamouroux. 
Langsdorf. 

La Peyrouse. 
La Pylaie. 
Ledebour. 
Lehmann. 
Lemaire. 
Lesquereux. 
Lessing. 
Lestiboudois. 
Léveille. 

L’ Heritier. 

L’ Herminier. 
Liebmann. 
Lightfoot. 
Lilijeblad. 
Lindberg. 
Lindblom. 
Lindenberg. 
Lindheimer. 
Lindley. 
Linneus. Also Z. 
C. Linnzus, son. 
Loddiges. 
Leefling. 


388 ABBREVIATIONS. 
Les. = Leselius. Naud. = 
Los. Loiscleur-Delongschamps. | Neck. 
Loud. Loudon. Nees or ! 
Lour. Loureiro. N.ab E. 
Ludw. Ludwig. T. Nees 
Lumn. Lumnitzer. Nestl. 
Lyngb. Lyngbye. Newb. 
Newm. 
Macf. Macfadyen. Negg. 
Macgil. MacGillivray. Nois. 
Magn. Magnol. [stein. | Nord. 
M. Bieb. Marschall von Bieber- | Jot. 
Marsh. Humphrey Marshall. Nutt. 
Mars. Marsili. Nyl. 
Mart. Martius. Nym. 
Mass. Massalongo. 
Mast. Masters. (Ed. 
Maxim. Maximowicz. (rst. 
Med. Medikus or Medicus. Oliv. 
aes Meisner or Meissner. Heut 
Meneq. Meneghini. Orph. 
Menz. Menzies. Ort. 
Mert. Mertens. Oudem. 
Metten. Mettenius. 
Mich. Micheli. P.deBeauc. 
eer André Michaux. - aa 
Michx. f. ¥. A. Michaux, son. Park. 
Midden. |Middendorff. Parl. 
Mill. Philip Miller. Pasq. 
Mill. J. John S. Mueller or Miller. | Pav. 
Miq. Miguel. Perl. 
Mirb. Mirbel. Pers. 
Mitch. John Mitchell. Philib. 
Mitt. Mitten. Planch. 
Moe. Mocino. G. Planch. 
Molk. Molkenboer. Pluk. 
Mont. C. Montagne. Plum. 
Moq. Moquin-Tandon. Peepp. 
Moric. Moricand. Poir. 
Moris. Morison. Poit. 
Morr. Morren. Poll. 
Moug. Mougeot. Post. 
Muell. Arg. J. Mueller of Argau. Pourr. 
F. Muell. Ferdinand Mueller. Pringsh. 
O. Muell. Otto Muellerof Denmark. | Pritz. 
Muhl. Muhlenberg. Putter. 
Munt Munting. 
Murr. J A. Murray. Rabenh. 
A. Murr Andrew Murray. Radlk. 
Raf: 
Nace. Naccari. Rasp. 
Neg. Negeli. Red. 


Naudin. 
Necker. 
C. F. Nees von Esenbeck. 


T.F.L. Nees von Esenbeck. 
Nestler. 
Newberry. 
Newman. 
Neeggerath. 
Noisette. 
Nordstedt. 
Notaris. 
Nuttall. 
Nylander. 
Nyman. 


Eder. 

(Ersted. 

Olivier. 

D. Oliver. 

A. or C. d’Orbigny. 
Orphanides. 
Ortega. 

Oudemans. 


Palisot de Beauvois. 
Pallas. 

Panzer. 
Parkinson. 
Parlatore. 
Pasquale. 

Pavon. 

Perleb. 

Persoon. 

Philibert. 

J. E. Planchon. 
Gustave Planchon. 
Plukenet. 

Plumier, Lat. Plumerius. 
Peeppig. 

Poiret. 

Poiteau. 

Pollich. 

Postels. 

Pourret. 
Pringsheim. 
Pritzel. 

Putterlich. 


Rabenhorst. 
Radlkofer. 
Rafinesque-Schmaltz. 
Raspail. 

Redoute. 


ABBREVIATIONS. 

Reich. = Reichard. Scop. = 
Reichenb. WH. G. L. Reichenbach. Seem. 
Reichenb. f. H. G. Reichenbach, son. Sendt. 
Reinw. Reinwardt. Seneb. 
Reiss. Reisseck. Ser. 
Retz. Retzius. Seub. 
Reut. Reuter. Sibth. 
Rich. L. C. Richard. Sieb. 
Rich. f. : : Sieb. 
ac pad } Achille Richard. Rae 
Liichards. John Richardson. Sow. 
Richt. Richter. Spenn. 
Ridd. Riddell. Spreng. 
Riv. Rivinus. Sternb. 
Reh. Rebling. Steud. 
Rem. J. J. Roemer. Stev. 
M. J. Rem.M. J. Roemer. Sull. 
eg Remer & Schultes. iat 

Sch. 
Rep. Reper. Targ: 
Rohrb. Rohrbach. Ten. 
Rostk. Rostkovius. Thoms. 
Rothr. Rothrock. Thuill. 
Rottb. Rottbell. Thunb. 
Rottl. Rottler. Thurb. 
Roum. Roumegere. Thurm. 
Roxb. Roxburgh. Tod. 
[oy. Royen. Torr. 
Rudb. Rudbeck. Torr. § 
Rupr. Ruprecht. Tourn. 

Tratt. 

Sace. Saccardo. Traut. 
Sadi. Sadler. Trev. 
St. Hil. A. Saint-Hilaire. Trin. 
Salish. Salisbury. Tuck. 
Salm-Dyck.Prince Jos. Salm-Riffer- | Turcz. 
Sauss. Saussure. [schied-Dyck. | Turn. 
Schimp. | Schimper. Turp. 
Schk. Schkuhr. 
Schlecht. Schlechtendal. Vaill. 
Schleich. Schleicher. Veill. 
Schomb. Schomburgh. Vauch. 
Schrad. Schrader. Vent. 
Schreb. Schreber. Vill. 
Schueb. Schuebeler. Vis. 
Schult. Schultes. “ttad. 
Schultz | C. H. Schultz, Bipontinus iv. 

Bip. (Zweibrucken) Vog. 
Schum. Schumacher. 
Schnitzel. Schnitzlein. Wahl. 
Schwagr. Schwegrichen. Wahist. 
Schwein. Schweinitz. Walds. 
Schweinf. Schweinfurth. Wall. 
Schwend. Schwendener. Wall. 


G 


~ 


Scopoli. 
Seemann. 
Sendtner. 
Senebier. 
Seringe. 
Seubert. 
Sibthorp. 
Sieber. 
Siebold. 
Solander. 
Sowerby. 
Spenner. 
Sprengel. 
Sternberg. 
Steudel. 
Steven. 
Sullivant. 
Swartz. 


Targioni-Tozetti. 


‘Tenore. 


Thomas Thomson. 
Thuillier. 
Thunberg. 
Thurber. 
Thurman. 

Todaro. 

Torrey. 


.Torrey & A. Gray. 


Tournefort. 
Trattinick. 
Trautvetter. 
Treviranus. 
Trinius. 
Tuckerman. 
Turezaninow. 
Turner. 
Turpin. 


Vaillant. 

Veillard or Vieillard. 
Vaucher. 

Ventenat. 

Villars, or Villar. 
Visiani. 

Vittadini. 

Viviani. 

T. Vogel. 


Wahlenberg. 
Wahlstedt. 
Waldstein. 
Wallich. 
Wallman. 


389° 


“390 


Wallr. = Wallroth. 


Walp. Walpers. 

Walt. Walter. 

Wang. Wangenheim. 
Warm. Warming. 
Wats. P. W. Watson. 


H.C.Wats.H. C. Watson. 


S. Wats. Sereno Watson. 
Web. W eber. 

Wedd. W eddell. 
Weinm. Weinmann. 
Welw. Welwitsch. 
Wender. Wenderoth. 
Wendl. Wendland. 
Wiks. Wikstrom. 


ABBREVIATIONS. 


Wildb. = Wildbrand. 


Willd. Willdenow. 
Willk. Willkomm. 
Wils. Wilson. 
Wimm. Wimmer. 
Wisliz. Wislizenus. 
With. Withering. 
Woode. W oodville. 
Wulf. Wulfen. 
Zanard. Zanardini. 
Zetterst. Zetterstedt. 
Zuce. Zuccarini. 
Zuccag.  Zuccagini. 


2. ABBREVIATIONS OF NAMES OF ORGANS AND TERMS 
¢ USED IN BOTANICAL WRITINGS. 


Est. Jéstate, in summer. 

“Est. Estivation. 

Alb. Albumen. 

Anth. Anther. 

Art. Artificial. 

Auct., Auctt. Auctorum, of authors. 

Aut. Autumnal. 

B. or Beat. Beatus, “the late,” re- 
cently deceased. 

Br. Bract. 

Cal. Calyx. 

Cel. Celeberrimus, or Very cele- 


brated. 
Cent. Centimetre. 
Cl. Clarissimus. 
Char. Character. 
Coll. Collection. 
Cor. Corolla. 
Cult. Cultivated. 


Decim. or Dec™. Decimetre. 

Deser. Description 

Diff. Differentiz, the distinguishing 
marks. 

Ed. Edition. 

Embr. Embryo. 

Ess. Essential, as Char. Ess. 

Excl. Excluding, or being excluded. 

Excl. Syn. Excluding the synonym 
or synonyms. 


Fam. Family. 

Fil. Filament of the stamen. 

Fl. Flower (flos); Flora, or some- 
times Floret, it flowers. 

Fem. Female plant, flower, &c. 

Fol. Folium, leaf. 

Fr. Fruit. 

Fructif. Fructification. 

Gen. Genus or Generic. 

Germ. Germen, Linnean name for 
ovary ; also Germination. 

H. Herbarium. 

Hab. Habitat, place of growth; 
sometimes for Habeo, I have. 

Herb. Herbarium. 

Hort. Hortus, garden. 

Hortul. Hortulanorum, of the gar- 
deners. 

Ic. Icon, a plate or figure. 

7/7. Mllustris, illustrious. 

Ined. Unpublished. 

Inf. Inferior. 

Infl. Inflorescence. 

Inv. Involucre. 

Lat. Lateral, or relating to width. 

Lin. Linea, a line (the 12th of an 
inch). 

Lit., Litt. In a letter or letters. 

| 1. c. Loco citato, in the place cited. 


ABBREVIATIONS. 391 


Musc. Male plant, flower, &c. | Sep. Sepal. 
Mill. or mm. Millimetre. Ser. Series. 
Mss. Manuscripts. | Sice. Siccatus or Siccus, dried or dry. 
Mus. Museum. Spec. or Sp. Species, or specimen. 
N. or No. Number. | Spont. Spontaneous. 
Nat. Natural. | Stam. Stamen or Staminate. 
Nom. Nomen, name. Sup. Superior. 
Obs. Observation. | Syn. Synonym or Synopsis. 
Ord. Order. | T. or Lab. Tabula, plate. 
Ov. Ovary. | ZY. Tomus, volume. 
p. Page, or sometimes Part. V. Volume: sometimes for Vel, or; 
Ped. Peduncle or Pedicel, or for sometimes Vide, see. 
Pedalis, a foot long or high. : Var. Variety. 


Peric. Pericarp. 
Perig. Perigonium. 


Pet. 
Pist. 


Place. Placenta. 


Poll. 
p- p- 


Prodr. or Prod. Prodromus. 


Rad. 


Ram. Ramus, branch. 
s. Seu, or Sive, Latin for or. 


Veg. Vegetation, cliaracters of 

Vern. Vernal. 

v.s. Visa sicca, or Vidi siceam. 

v.v. Visa viva, or Vidi vivam; the 
first indicating that a dried speci- 
men of the piant, the second that 
the living plant has been exam- 
ined. 

» Ss. c. and v. &. &., indicates that 
the dried specimen was cultivated 
(c) or spontaneous (s). 


Petal or Petiole. 
Pistil. 


Pollicaris, an inch long. 
Pro parte, in part. 


I 


Radix, root; or Radical. 


Sect. Section. v.v. c. and v. v. s., that the living 
Segm. Segment. plant seen was cultivated (c) or 
Sem. Semen, seed. spontaneous (s). 

SIGNS. 

1. SIGNS USED BY LINNAUS. 

© _ An annual plant. 
g A biennial. 
2 A perennial. 
5 A tree or shrub. 
* Affixed to a reference, means that a good description will be found there. 
+ Indicates an obscure or doubtful species. 


‘) 
@ 


2. SIGNS USED BY DECANDOLLE AND LATER 
WRITERS. 


A monocarpic plant, 7. e. which dies after once flowering and fruiting, 
either annual or biennial, or of longer duration. 

Annual. 

Biennial. 

Monocarpic perennial, such as Agave. 


392 SIGNS. 


Perennial herb. 

Suffruter, an undershrub. 

Frutex, a shrub. 

Arbuscula, a tree-like shrub of ten to twenty-five feet in height. 

Arbor, a tree. 

A climbing plant. 

An evergreen. 

Male plant or flower. 

Female plant or flower. 

Hermaplirodite plant or flower. 

Indefinitely numerous, e. g. 00-andra, polyandrous. 

A sign of doubt. “ Thalictrum? Japonicum,’ doubts if the plant is 
really a Thalictrum. “ Thalictrum Japonicum, Thunb.?” doubts if the 
plant in hand is truly the species of Thunberg. Thalictrum Japoni- 
cum, Thunb., Willd.? doubts whether Willdenow’s 7. Japonicum is 
really that of Thunberg. 

A sign of certainty. As “ Thalictrum anemonoides, Michx.! FI]. Bor. Am. 
p. 822,” as used by DeCandolle, affirms that he nas 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. 

°//’ 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. 

©]! Cotyledons incumbent on the radicle. 


~ Bw BPD D UMULtEFR 


GLOSSARY, 


OR 


DICTIONARY OF BOTANICAL TERMS, ENGLISH 
AND LATIN, 


COMBINED WITH AN INDEX. 


Tuts 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 Phzenogamous 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 umnecessary 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 | Abruptly pinnate. Pinnate without a 


words of Greek derivation, signifies terminal leaflet or appendage; 101. 
the absence of the organ mentioned; | <Acanthdcladous (-us). Having spiny 
as, Apetalous, destitute of petals; branches. 


Aphyllous, leafless. In words be- | Acanthéphorous (-us). Spine-bearing. 
ginning with a vowel, this prefix is | Acaulescent (-ens). Stemless, or appar- 


changed to an; as, Ananthous, flow- ently so, with no proper caulis; 45. 

erless ; Anantherous, antherless. Acaulis. Stemless; same as Acaulescent. 
Abbreviations, 385. Accessory. Something additionai, or of 
Aberrant. Wandering, applied to spe- the nature of appendage. 

cies, genera, &c., which differ in some Accessory Buds, 44. 

respect from the usual or normal char- | Accessory Fruits, 300. 

acter of the group they belong to. Accrescent (-ens). Increasing in size 
Abnormal (Abnormis). Differing from with age, as often occurs with the 

the normal or usual structure. calyx after flowering. 
Aboriginal. Strictly native; indigenous. Accrete (-us). Grown together, or con- 
Abortion (Abortus). Imperfect develop- solidated with some contiguous body. 


ment or non-development of anorgan; | Accumbent (-ens). Lying against an- 
179, 187. | other body. 
Abortive (-ivus). Defective or barren. Accumbent Cotyledons. With edges 
Abrupt (Abruptus). Terminating sud- | against the radicle: 313. 
denly ; the opposite of tapering. Acephalous (-us). Headless. 


5904 


Acerose (-dsus). Needle-shaped, like the 
leaves of Pines. 

Acetabuliform (-ormis). In the form of 
a shallow open cup or saucer. 

Achen.um or Achenium. A small, dry 
and hard, one-celled, one-seeded, inde- 
hiscent fruit; strictly one of a single 
and free carpel; but extended tu simi- 
lar ones of more than one carpel, and 


also with adnate calyx; 294. (Ache- 
nium is etymologically the proper 
orthography; but achenium is De- 


coming the commoner form.) 

Achenocarp (-arpium). General name 
of a dry and indehiscent fruit ; 292. 

Achenodium. Such a double achenium 
as that of Umbellifere; a Cremo- 
carp. 

Achlamydeous (-eus). 
anth; 191. 

Acicula. <A bristle. 

Acicular (-aris). Bristle-shaped, or slen- 
der needle-shaped. 

Acinaciform (-ormis). Seymitar-shaped ; 
curved with rounded point, thicker on 
the straighter edge than on the con- 
vex edge. 

Acinosus. Like grapes or grape-seed. 

Acinus. Classically a berry, particu- 
larly a grape, or its stony seed, or a 
bunch of berries; now sometimes ap- 
plied to the separate carpels of an 
aggregate baccate fruit, or to the con- 
tained stone or seed; 297. 

Acorn. Fruit of the Oak. 

Acotylédon, p|. Acotyledons, Acotylédones. 
A plant or plants destitute of coty- 
ledon, or 

Acotyledonous (-eus). Without cotyle- 
dons; as the embryo of Cusecuta; 26, 
38. Mostly applied, as by Jussieu. 
to plants which have no proper seed 
nor embryo, and therefore no cotyle- 
don; 339. 

Acramphibry1. Plants producing side as 
well as terminal buds or growths ; 341 

Acrobrya. Plants growing from apex 

only; 341. 

-Acrogen (Acrdégene). Name of class of 
plants which in growth are said to be 

Acroyenous. Growing from the apex 
or by terminal buds only. 

Acrosércum. Desyaux’s name for a 
berry from an ovary with adnate 
calyx. 

Acrospira. An old name of the plu- 
mule of a grain in germination. 

Aculeate (-eatus). Prickly; beset with 
aculei. 


Destitute of peri- 


GLOSSARY. 


Aculeosus. Abounding with prickles. 

Aculeolate (-atus). Beset with diminu- 
tive prickles, or 

Aculeoli. Diminutive of aculei. 

Aculeus. A prickle; a pointed small 
excrescence of the bark. 

Acumen. <A tapering point. 

Acuminate (-atus). Ending in a tapering 
point; 96. 

Acute (Acutus). Terminating in an acute 
angle; 97. 

Acropetal. Developing from below up- 
ward, or from base toward apex. 

Actinomorphous (-us). Capable of bi- 
section through two or more planes 
into similar halves, as is a regular 
symmetrical flower; 175. 

Acutiisculus. Somewhat acute; acutish. 

Adelphous (-us, Adelphi, brothers). Sta- 
mens with cealescent or clustered fila- 
ments are monadelphous, diadelphous, 
&c., according to the number of Adel- 
phia or brotherhoods. 

Aden. Greek fur gland, is compounded 
with Greek words with this meaning; 
as, Adendphorus, gland-bearing; -Ad- 
enophyllus, leaves bearing glands, &c. 

Adylutinate (-atus). Same as accrete. 

Adherent (Adherens). Generally same 
as adnate; may refer to adhesions not 
congenital. 

Adnate (-atus). Congenitally united to ; 
as the calyx-tube of the gooseberry to 
the ovary; 182. Adnate anther is one 
seemingly borne on the outer or inner 
face of the filament ; 7. e. extrorsely or 
introrsely fixed by its whole length to 
the connective; 253. 

Adnation. The state of being adnate; 
179; 181. 

Adpressus. Latin of appressed. 

Adscendens. Latin of ascending. 

Adsurgens. Latin of assurgent. 

Adventitious, Adventive. That which has - 
come from abroad or as astranger; as 
a plant lately or by chance introduced 
from another country. 

Adventitious Buds; 45. 

Ayquilateralis. Equilateral, equal-sided. 

Aqualiflérus. When all the flowers of 
the same head or ciuster are alike in 
form as well as character. 

Ayudlis, Aquans. Equal; equalling. 

Aérial roots, &e. ; 33. 


Aérophytes. Air-plants; 35. 
Eruginosus. Verdiaris-colored. 


stival (-dlis). Relating to summer. 
Estivation (-io). The disposition of the 
parts of a flower in the bud; 132. 


GLOSSARY. 


ZEterio. A form of aggregate fruit; 300. 

Athevgamia, Ethogamous, 340. 

Ajinity. True and near relationship; 
327, 330. 

Ayamous or Agamic. Destitute of sexes. 


aciiomenate Heaped or crowded 
pis ee eur) jin a dense cluster, 
La “but not cohering. 


Aggregate Fruits. Those formed of 
aggregate carpels of the same flower; 
248, 301. 

Agrestis. Growing in fields. 

Air-plunts. Plants unconnected with the 
ground; 35. > 

Akene, Akenium. See Achzenium. 

Ala (pl. ale). A wing. Also the side 
petals of a papilionaceous corolla; 
185. Has also been used in the sense 
of axilla. 

Alabastrum. <A flower-bud; 40. 

Alar (Alaris). From ala in the sense of 
axilla, therefore wxillary or in the forks. 

Alate (-atus). Winged. 

Albescens, Albicans. Whitened, whitish, 
or hoary. 

Albumen of the seed. Any deposit of 
Nutritive material within the seed- 
ccats, and not in the embryo; 14, 309. 

Albuminous or Albuminose ( Albuminvosus). 
Said of seeds provided with albumen; 
13, 309. 

Albirnum. Sapwood; the newer wood 
of an exogenous stem; 80. 

Albus. White. 

Allagostémonous. With stamens alterna- 
tively inserted on the torus and on the 


petals. 

Alliaceous (-eus). Having the smell of 
garlic. 

Alliance. Synonym of Cohort; 326. 


Allégamy. Fecundation of the ovules of 
a flower by other than its own pollen ; 
cross fertilization, 216. 

Alp:strine (Alpestris). Growing on 
mountains below an alpine region 
or one unwooded from cold. 

Alpine (-inus). Growing on the higher 
parts of the Alps, or (by extension of 
meaning) on other mountains above 
the limits of trees. 

Alternate (Altérnus). One after an- 
other; as of leaves placed singly 
instead of in pairs (opposite) or in 
whorls. Also, standing before inter- 
vals; as stamens alternate with petals 
instead of before them; 6, 119. 

Alternative (-ivus). In eestivation, with 
an inner whorl alternating with an 
outer one; 134, 136. 


395 


Alvéolate(-atus). Honeycombed ; having 
deep angular cavities (Alvéoli) sepa- 
rated by thin partitions, as the recep- 
tacle of cotton-thistle. 

Ambitus. The ray or circumference of 
a head, &c. 

Ament {Amentum). A catkin, or pe- 
culiar scaly spike; 150. 

Amentuceous (-cus). Bearing catkins, or 
catkin-like. 

Amorphous (-us). 
definite form. 

Amphanthium. One of the (needless) 
names coined for a dilated receptacle 
of inflorescence. 

Amphibrya. Equivalent to Monocotyle- 
dones; 341. 

Amphicarpous (-us). 
kinds of fruit. 

Amplhigamous Cryptogams, 340. 

Amphigastria. Peculiar leaves (of He- 

_ paticze) imitating stipules. 

Amphisarca. A hard-rinded berry, or 
fruit succulent within and woody or 
crustaceous without, as a calibash. 

Amphispermium. Link’s name for a 
one-seeded pericarp which is con- 
formed to the seed; an akene. 

Amplitropous (-us), wrongly Amphi- 
tropal. Turned both ways; applied 
to an ovule with hilum intermediate 
between micropyle and chalaza; 279. 

Amphora. A pitcher; and the lower 
part of a pyxis. 

Amplectens, Amplexans, Amplexus. Em- 
bracing, clasping. 

Amplezicaul (-aulis). Clasping a stem, 
as does the base of certain leaves. 

Ampliate (-atus). Enlarged or dilated. 

Ampulla. A bladder or flask-shaped 
organ, as of Utricularia. 

Ampullaceous (-us), or Ampulleform. 
In the form of a bladder or short flask. 

Amylaceous (-eus). Resembling or com- 
posed of starch, or Amylum. 

Amyloid. Analogous to starch. 

Analogy (Analogia). Likeness in cer- 
tain respects. As distinguished from 
affinity, it means resemblance in cer- 
tain respects on/y, 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 analogues ; t. € , are 
analogous organs, but are not homo- 
logues ; for the first answers to a leaf, 
the second to stipules, and the third 


Shapeless; of in- 


Producing two 


396 


toastem. 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. 

Anandrous. Destitute of stamens. 

Anantherous (-us). Destitute of anthers. 

Aninthous (-us). Flowerless. 

Anastomosis. The connection of veins, 
&c., by eress-veins, forming reticu- 
lation. 

Andtropous (-us), wrongly Anatropal. 
The reversed ovule, with micropyle 
close by the side of the hilum, and 
chalaza at the opposite end; 279. 

Anceps, Engl. <Ancipital. Botanically 
always used in the sense of two- 
edged. 

Ander, andra, andrum. 
pounds, the male. 
Andro-diecious. With flowers on one 
plant hermaphrodite, and on another 

staminate only; 191. 

Andrecium. The stamens of a flower 
collectively ; 165, 249. 

Androyynous (-us). Said of an inflores- 
cence composed of both male and 
female flowers. 

Androphore (Andréphorum). A sup- 
port or column on which stamens are 
raised. 

Androus. See Ander. 

Aneméphilous. 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. 

Angiocarpous (-us). When a fiuit is 
covered by some envelope. 


In Greek com- 


Angiospermia. A Linnzan artificial 
order; 337. 

Angiospermous, Angiosperme, Anyio- 
sperms. Plants with seeds borne ina 


pericarp; 259. 
Angular Divergence of leaves; 123. 
Anisémerous (-us). Unequal in number 
in the different circles of the flower; 
unsymunetrical. 
Anisopélalous (-us). With unequal petals. 
Anisuphyllous (-us). | Unequal-leaved; 
i. €., the two leaves of a pair unequal 
Anisostémonous (-us) When the sta- 
mens are not of the number of the 
petals. 
Annotinus. 
growths. 
Annual (Annuus) 
duration; 30. 


A year old, or in yearly 


Of only one year’s 


GLOSSARY. 


Annular (-aris). In the form of a ring; 
or marked transversely by rings. 
The latter more properly. 

Annultte (-atus). Marked with rings. 

Annulus. A ring, such as that with 
which the sporangia of some Ferns 
aud Mosses are furnished. 

Anophytes (Anophyta) Name of group 
comprising Mosses, &e. 

Antepositio: Same as Superposition; 
179, 195. 

Anterior, as to position, denotes the front 
side, or averse from the axis of inflo- 
rescence; 160. 

Anthela. A deliquescent and paniculate 
cyme, with median ramification, and 
the lateral axes overtopping the 
central, as in Juncus tenuis, &e. 
May be either a Drepanium or a Rhi- 
pidium. 

Authemy, Anthemia. 
of any kind; 144. 

Anther (Anthera). The polliniferous 
part of a stamen; 165, 251. : 

Antheridium. An analogue of the an- 
ther in Cryptogams. 

Antheriferous (-us). Anther-bearing. 

Anthesis. The time at which a flower 
is perfected and opens; or the act of 
expansion of a flower. 

Anthocarpous(-us), Anthocarpuum. Fruits 
in which some organ exterior to the 
pericarp is concerned ; 300. 

Anthoclinium. Name of a receptacle of 
inflorescence, such as that of Com- 
posite. 

Anthodium. A name for the head of 
flowers (or so-called compound flowers) 
of Composite; 147. 

Antholysis. A retrograde metamor- 
phosis of a flower, in which normally 
combined parts are separated. 

Anthophore (Anthéphorum). The stipe 
when developed between calyx and 
corolla; 212. 

Anthus or Anthos. 
compounds. 

Anticous (Anticus). 
253. 

Antitropous (-us), less properly Antitro- 
pal. Said of an embryo with radicle 
pointing to the end of the seed oppo- 
site the hilum; 312. 

Antrorse. Directed upward or forward. 

Apetalous (-us). Having no petals; 190. 

Apex. Besides its ordinary meaning, 
the top of a thing, it was once the 
technical name of an anther; 166. 

Aphyllous (-us). Leafless. 


A flower-cluster 


A flower, in Greek 


Facing anteriorly ; 


GLOSSARY. 


Apical (-alis). Relating to the apex or tip. 

pices. The name for anthers anterior 
to Ludwig and Linnzus; 166. 

Apiculate (-us). Ending in a short 
pointed tip or apicula. 

Apocarpous (-us). When carpels of a 
gyucecium are separate; 261, 262. 

Apéphysis. An enlargement or swelling 
of the surface of an organ at some par- 
ticular part. 

Apothecia. ‘The ‘shields ”’ or fructify- 
ing disks of Lichenes. 

Apotropous (-us). Said of an anatropous 
ovule which when pendulous has 
rhaphe averse ; 282. 

Appendage, Appendix. Any superadded 
or subsidiary part. 

Appendiculate (-atus). Furnished with a 
small appendage (Appendiculum), or 
with any appendage. 

Appositus. Placed side by side. 

Appressed (Lat. Adpressus). Lying flat 
against or together for the whole 


length. 
, . . . 
Apricus. Growing in dry sunny places. 


Apterous (-us). Wingless; not alate. 
Aquatic (-icus). Living in water. 
Aquatilis. Living under water. 
Arachnoid (-oideus). Cobwebby ; com- 
posed of slender entangled hairs. 

Araneose (-osus), Araneus. Like spider- 
web; same as Arachnoid. 

Arbor. <A tree; 50. 

Arboreous ( Arboreus). 
lating to a tree. 

Arborescent (-ens). Tree-like; approach- 
ing the size of a tree. 

Arboiétum, also Arbustum. <A_ place 
where trees are grown; an arranged 
collection of trees. 

Arbiscula. A small shrub of tree-like 
growth or form. 

Arbuscularis. Ramified like a little tree. 

Archegonium. The spore-case of mosses, 

_&c., in an early state. 

Arcuate (-atus). Moderately curved, as 
if bent like a bow. 

Areéola, pl. Areole. Spaces marked out 
on a surface, as by the reticulation of 
veins, &e. 

Aréolate (-atus). Marked with areole. 

Arenosus, Arenarius. Growing in sand 
or sandy places. 

Argentate (-atus, Argénteus). Silvery, 
or shining white with a tinge of gray. 

Argillosus. Growing in clayey soil. 

Argos. Greek for pure white, used in 
compounds; as, aryophyllus, white- 
leaved. 


Tree-like, or re- 


397 


Argitus. Sharp-toothed; said of the 
serration of leaves. 

Argyros. Greek for silvery; used in 
compounds; as, argyrophyllus, silvery- 
leaved. 

Arhizal ( Arhizus). Rootless. 

Avillute (-atus). Having an arillus. 

Avil, Arillus. Aw extraneous or late- 
formed seed-coat or covering, or an 
appendage growing from or about the 
hiluin of a seed; 308. 

Arillifurm (-ormis). In the form of an. 
arillus. 

Arillode, Arillodium. <A false arillus, 
or one which does not originate from 
or below the hilum, but from the 
micropyle or rhaphe; 309. 


Arista. An awn. 

Avistate (-atus). Awned; bearing an 
arista. 

Aristulate (-atus). Bearing a diminu- 
tive awn. 


Arrect (Arrectus). 
upright position. 

Arrow-shaped, Arrow-headed. Same as 
Sagittate; 96. 

Articulated (-utus). Jointed, or having 
the appearance of a joint or articula- 
tion ( Articulus). As of the word joint 
itself, the context must show whether 
the articulations mean the portions 
which are connected by a joint, or the 
place of connection. 

Artyicial Classification, 331. 

Ascending ( Adscendens). Rising upward. 
Sometimes used for directed upward, 
as when the stem is termed the As- 
cending Axis (11); more commonly 
denotes curving or rising obliquely 
upward; 53. 

Ascidium. A pitcher-shaped or flask- 
shaped organ or appendage; 111. 

Ascus. A sac; a kind of spore-cases, as 
in certain Fungi and Lichenes. 

Asparagi. A name for Turiones, or any 
scaly shoots from underground, as 
those of Asparagus. 

Aspergilliform (-ormis). Brush-shaped, 
i.e. like the aspergillum, or brush 
used to sprinkle holy water; made up 
of numerous spreading hairs, &c., in 
a tuft. as the stigmas of Grasses. 

Asperous (Asper). Rough to the touch. 

Assimilation. The action or process by 
which extraneous matter or crude food 
is converted into vegetable matter. 

Assurgent (Adsurgens). Rising or cury- 
ing upward; 53. 

Astichous (-us). Not in rows. 


Brought into an 


398 

<Astomous (-us). Without a stoma or 
mouth. 

Atavism (-mus). Ancestral resemblance. 

Ater. Pure black. 

Athera. Greek for Arista or Awn. 

Atratus. Blackened or turning black. 

Atrepous (-us), wrongly Atropal. Not 
turned; applied to an ovule the same 
as orthotropous ; 277. 

Attenuate (-atus). Slenderly tapering 
or narrow. 

Auctus. Same as accrescent; enlarged 
after flowering ; augmented by an ad- 
dition. 

Augmentation. Increase beyond the 
normal number; 179, 200. 

Aurantiacus. Orange-colored. 

Auratus, Aureus. Golden-colored, or 
yellow with golden lustre. 

Auricle (Auricula). An ear or ear- 
shaped appendage. 

Auriculate (-atus). Furnished with an 
auricle; 96. 

Autocarpous. A fruit consisting of peri- 
carp alone, having no adnate parts. 
Autogomy. Close-fertilization, the fe- 
cundation of a flower by its own pol- 

len; 215, 216. 

Avenius. Veinless. 

Auwl-shaped. Narrow, terete or some- 
what so, and attenuate from a broader 
base to a slender or rigid point. 

Awn. A bristle-shaped appendage, such 
as the beard of Rye and Barley. 

Awned. Furnished with an awn. 

Azil (Azilla). The angle formed on the 
upper side of the attachment of a leaf 
‘with the stem, or the point just above 
this attachment; 6. 

Axillary (-dris). In or relating to an 
axil; 7. 

Azile, Azial ( Azilis). 
longing to the axis. 

Azis. The stem; the central part or 
longitudinal support on which organs 
or parts are arranged ; the central line 
of any body. 


Relating or be- 


Bacca. A berry; 299. 

Baccate (-atus). Berry-like; pulpy 
throughout. 

Baccetum. An aggregation of berries 


in one flower; 300. 

Badius. Chestnut-brown. 

Balausta. Name applied to the fruit 
of the Pomegranate, with firm rind, 
crowned with the lobes of an adnate 
calyx, baccate within, and many- 
seeded. 


GLOSSARY. 


Banner. The vexillum, standard, or 
upper petal of a papilionaceous co- 
rolla; 184. 

Barb. A bristle or stout hair, which is 
hooked or double-hooked, or retrorsely 
appendaged at the tip. 

Barba. Beard. 

Barbate (-atus). Bearded; beset with 
long and weak hairs. 

Barbellate (-atus). Beset with shorter 
and stiffer hairs or barbelle. 

Barbellulate (-atus). Diminutive of the 
preceding. : 

Bark. The rind or cortical portion of a 
stem, especially of an exogen; 76. 

Basal ( Basilaris). Relating to the base. 

Basé@!-nerved. With nerves all from the 
base of the leaf; 92. 

Base (Basis). The extremity by which 
an organ is attached to its support. 
Basidia. Cells of the fructification of 
Mushrooms which bear the spores. 
Buasificed (-us). Attached by the base 

or lower end; 253. 

Basigynium. Synonym of Carpophore 
or Thecaphore. 

Basinerved (-ius). When the ribs pro- 
ceed from the base of a leaf. 

Busipetal. Developing from apex to- 
ward the base. 

Bast, or Bass. Inner fibrous bark; 77. 

Bast-cells. The es-ential components of 
bast; long and flexible butthick-walled 
attenuated cells; 77. 

Beak. A narrowed or prolonged tip. 

Beaked. Ending in a beak. 

Bell-shaped. Same as Campanulate ; 249. 

Berried. Baccate. 

Berry. A fruit, the whole pericarp of 
which is fleshy or pulpy; 299. 

Bi- or Bis. As a prefix to Latin words 
(Greek words have Di-), two, twice, 
or doubly. 

Biacuminate (-atus). _Two-pointed, as 
malpighiaceous hairs, fixed by the 
middle and tapering to each end. 

Biarticulate (-atus). Two-jointed. 

Biauriculate (-atus). Two-auricled. 

Bibracteate ‘-atus). With two bracts. 

Bibracteolate (-atus). With two bract- 
lets. 

Bicallose (-osus). 

Bicarinate (-atus). 


With two callosities. 
Two-keeled. 


Biceps. With two supports or stalks, 
Bicipital. ) or two-headed. 
Bicolor. Two colored. 


Biconjugate (-atus). Twice paired. 
Bicornis. Two-horned. 
Bicurnute. Saie as preceding. 


GLOSSARY. 


Bicruris. Two-legged, or with two sup- | ABipinnate (-atus). 


ports. 

Bidentate (-atus). Having two teeth. 
(Not doubly dentate.) 

Biluus. Lasting two days only. 

Biennial ( Biennis). Of two years’ dura- 
tion; 31. 

Bifurious (-ius). 
vertical rows. 

Biferus. Double-bearing; fruiting twice 
a year. 

' Bifid (-idus). Two-cleft, to the middle 
or thereabout. 

Bifldrous (-us). Fwo-flowered. 

Bifoliate. Two-leaved. 

Bifoliolate. Of two leaflets. 

Biferate (-atus). Having two open- 
ings. 

Biformis. Two-formed; in two shapes. 

Bifvrons. With two faces or aspects. 

Bifireate (-atus). Two-forked; i.e. of 
two prongs or forks. But it may 
mean bis furcatus; i.e., forked and 
again forked. 

Bigeminate (-atus). 
as Biconjugate. 

Biyener. The offspring of a cross be- 
tween two generically different plants. 

Bijugate (Bijugus). Two-paired, as a 
pinnate leaf of two juga or pairs of 
leaflets. 

Bilabiate (-atus). Two-lipped; 247. 

Bilamellate (-atus), or Bilamellar. Of 
two plates or lamellae. 

Bilobed ( Bilobus), or Bildbate. Of two 

_ lobes, or cleft into two segments. 

Bilocellate. Divided into two locelli; 
263. 

Bilocular (-arvis). Two-celled. 

Bimestris. Lasting two months. 

Bimus. Lasting two years; two years 
old. 

Binary (-arius). 
members; 176. 

Binate (-atus). In pairs or twos. 

Bini. Twin, or two together. 

Binodal ( Binddis). Having two nodes. 

Binomial Nomenclature, 346. 

Bivlogy. “The natural history of plants 
and animals, 7. e. of living things; 1. 

Bipalmate (-atus). Twice palmately com- 
pound. 

Biparous. Bearing two; as a cyme of 
two rays or axes; 152, 155. 

Bipartible (-ibilis). Capable of division 
into two similar parts. 

Bipartite (-itus). Divided almost into 
two pieces; two-parted. 

Bipes. Same as Bicruris. 


Two-ranked; in two 


Twice twin; same 


Consisting of two 


399 


Doubly or twice 
pinnate; 103. 

Bipinnatifid (-idus). 
pinnatifid; 100. 

Bipinnatisect (-us). Twice pinnately di- 
vided. 

Biplicate (-atus). Twice folded or plaited. 

Biporose (-osus). Opening by two pores. 

Biradiate (-atus). Of two rays. 

Birimose (-osus). Opening by two slits. 

Bisected (-us). Comyletely divided into 
two parts; 99. 

Biseptate (-atus). With two partitions. 

Biserial (-ialis), or Biseriate (-iatus). In 
two series, one above the other. 

Biserrate (-atus). When serratures are 
again serrate: doubly serrate. 

Bisexual. Having both stamens and 
pistil; hermaphrodite; 191. 

Bisilcute (-atus, Bisulcus). Two-grooved; 
having two furrows. 

Bitérnate (-atus). Twice ternate. 

Bladdery. Thin and inflated. 

Blade. The lamina, limb, or expanded 
portion of a leaf, &e.; 85, 245. 

Blastéma. The budding or sprouting 
part or point. First used for the axis 
of an embryo; now used for the ini- 
tial growth out of which any organ 
or part of an organ is developed. 

Bloom. Besides its use as equivalent to 
blossom, it denotes the white powdery 
and glaucous covering of the surface 
of many fruits and leaves, of a waxy 
nature. 

Boat-shaped. Of the shape of a boat, of 
the deeper sort, with or without a keel. 

Bostrychoidal. Having the form or char- 
acter of a ringlet, or Bostryx: 157. 


Twice. or doubly 


Bostryz. An uniparous helicoid cvme; 
156. 
Bothrénchyma. Tissue of plants com- 


posed of dotted or pitted ducts. 
Botry-cymose. Racemes or any botrvose 
clusters cymosely aggregated ; 159. 
Botryose (-osus), Botryoidal. Of the ra- 
cemose type; 144. 145, 146, 153. 
Botrys. The equivalent of Raceme ; 146. 
Botuliformis. Sausage-shaped. 
Biachiate (-idtus). With spreading 
arms, as branches (especially opposite 
and decussate) widely diverging. 
Brachys. Greek for short, and used in 
compounds; as, Brachypodus, short- 
stalked. 
Bract, Bractea. The leaves (more or 
less modified) of a flower-cluster; 118, 
141. 


Bracteate (-eatus). Having bracts. 


400 


Bractéola, Bracteole. See Bractlet. 

Bracteolate(-atus). Having bractlets. 

Bractlet. A bract of the ultimate grade, 
as one inserted ona pedicei or ultimate 
flower-stalk, instead of subtending it; 
141, 142, 160. 

Bracteose (-osus). 
spicuous bracts. 

Branches. Secondary axes, or divi- 
sions of an axis; 47. 

Branchlets. Ultimate branches or divi- 
sions of an axis; 47. 

Breathing-pores. See Stomata, 89. 

Bristle. A stiff hair, or any slender 
body or outgrowth which may be 
likened to a hog’s bristle. 

Bristly. Beset with bristles. 

Brunneus. Deep brown. 

Brush-shaped. See Aspergilliform. 

Bryology. The botany of Mosses. 

Bud. The undeveloped state of a stem 
or branch, with or without leaves; 
6, 40. 

Bud-scales. The teguments of a bud; 40. 

Bulb ( Bulbus). A \eaf-bud (commonly 
subterranean) with fleshy scales 
eats; 43, 62. 

Bivbiceps. A stem with bulbous base. 

Bulbiferous (-us). Bulb-bearing. 

Bulbiilus, Builbulus. Diminutive bulb. 
Same as 

Bulblet. A small bulb, especially such as 
is produced in the air, in the axil of or- 
dinary leaves, or upon them; 63. 

Bulbodium. A synonym of Corm, the 
“solid bulb.” 

Bulbo-tuber. Synonym of Corm. 

Bulbous, Bulbosus. Having bulbs or the 
structure of a bulb. 

Bullate (-atus). Said of a puckered sur- 
face (as if blistered), thrown into por- 
tions which are convex and projecting 
on one side and concave on the other. 
Also used in specific names, in its 
more literal sense for inflated. 

Bursicula. A small pouch (bursa); 
such as that which encloses the disk 
or gland of the caudicle of the pollin- 
ium of an Orchis. 

Bursiculatus. Furnished with a bursi- 
cula or pouch. 

Byssaceous (-eus). Composed of fine 
threads, like dyssus or fine flax. 


Full of, or with con- 


Cadicous (-us). Dropping off very early, 
as the calyx of a Poppy at the time 
of expansion; 243. 

Ceruleus. Sky blue. or pure blue. 


or 


} 


GLOSSARY. 


Cesius. Lavender-color; pale green with 
whitish or gray. 

Calathidium, Calathis. Literally a bas- 
ket; aname for the head of flowers (or 
better for the involucre only) of Com- 
positee 

Calathiform (-ormis). Cup-shaped ; of 
somewhat hemispherical outline. 

Calcar. A spur; mostly used for the nec- 
tariferous one of a calyx or corolla. 

Calcarate (-dtus). Furnished or pro- 
duced into a spur. 

Calceolate (-atus), or  Calceiformis. 
Shaped like a slipper or shoe. 

Callose (-osus). Bearing  callosities 
(calli), or hard protuberances. 

Calvus. Bald, as an akene without 
pappus. 

Calycanthemy. Name of the monstros- 
ity in which the calyx imitates an 
exterior corolla: 174. 

Calyciflorous (Calyciflore), 340. 

Calycine (Calycinus). Relating to calyx. 

Jalyculate (-atus). Bearing bracts next 
the calyx which imitate an external 
or accessory calyx. 

Calyculus. An involucre or involucel 
imitating an additional calyx. 

Calyptra. The hood or veil of the 
spore-case of a Moss; or some cover- 
ing body like it. 

Calyptrate (-atus). Furnished with a 
calyptra, or something like it. 

Calyptriform (-ormis). Calyptra-shaped ; 
as the calyx of Eschscholtzia. 

Calyz. The flower-cup, the exterior 
perianth ; 164. 

Camara and its diminutive Camérula 
(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 layer ; 78. 

Campanulate (-atus). Bell-shaped ; elon- 
gated cup-shaped or shorter, and broad 
from the base; 249. 

Campaniformis. Same as Campanulate. 

Campylosp:rmous (-us). Curved-seeded. 
Said of seed-like fruits or carpels, as 
those of some Umbellifere, in which 
the contained seed is involute by the 
lateral edges, so as to produce a longi- 
tudinal furrow on the ventral face. 

Campylotropous (-us), or less correctly 
Campylotropal, or Campulitropous. 
An ovule or seed which is curved 
in its formation so as to bring the 


GLOSSARY. 


micropyle or true apex down near to 
the hilum; 279. 
Canalicaulate (-atus). Channelled, or 
with a longitudinal groove. 
Cancellate (-atus). Latticed; resembling 
lattice-work. 


Candidus. Pure white. 

Canescens. Hoary, usually with gray 
pubescence. 

Canus. Gray-white; whiter than the 


preceding. 

So slender that it 

may be compared 

with the hairs of 
animals. 

Capitate (-atus). WHead-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. 

Capreolate (-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. 

Capsuliferous. Capsule-bearing. 

Carcvulus. 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. 

Carinate (-alus). Keeled. 

Cariopsis or Caryopsis. A grain; a seed- 
like fruit with thin pericarp adnate to 
the contained seed ; 295. 

Carneus. Ylesh-colored, very pale red. 

Caro. Flesh, as the pulp of a melon, or 
the fleshy part of a drupe. 

Carpadilium. Synonym of Cremocarp. 

Carpel, Carp/llum. A simple pistil, or 
an element of a compound pistil, an- 
swering to one leaf; 167, 260. 

Carpid, Carpidium. Synonym of carpel. 

Carpology. The botany of fruits. 

Carpophore (Carpéphorum). A portion 
of receptacle prolonged between the 
carpels; 212. 

Carpophyll (Carpophyllum). Literally 
fruit-leaf; svnonym of Carpel; 260. 
Cartilaginous or Cartilagineous (-eus). 
Of the texture of cartilage or gristle ; 

firm and tough. 


Capillaceous (-eus). 
Capillary (-aris). F 


401 


Caruncle (Carincula). 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. 

Caryopsis. See Cariopsis. 

Cassideus. Helmet-shaped. 

Cassus. Empty, as an anther contain- 
ing no pollen. 

Castrate (-atus). Said of a stamen 
which wants the anther. 

Catapetalous (-us). Where petals are 
united only by cohesion with united 
stamens, as in Mallow. 

Catuphylla. Answers to the German 
‘* Niederblatter,’’ or under-leaves, 
those at the beginning of a growth, 
cotyledons, bud-seales, 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, &e. 
Caulescent (-ens). 

stem. 

Cailicle (Cauliculus). The initial stem 
in an embryo, generally named the 
Radicle; 10. 

Cauline (-inus). Belonging to the stem. 

Caulis. Greek form Caulon. The stem 
of a plant. 

Caulocaipic or Caulocarpous. Applied 
to plants which live to flower and 
fructify more than once or indefi- 
nitely. 

Cauléme, Cauloma. 
plant. 

Cephalanthium. One of the names of 
the head or capitulum in Composite ; 
148. 

Cell ( Cellula). 


Having an obvious 


The stem-part of a 


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 
seeds; 262. 

Cellular Plants, Cellulares, 340. 

Céllule (-ula). Diminutive of cell; of 


402 


the same meaning as Cell in vegetable 
anatomy; 28. 

Cellulose. The material, chemically con- 
sidered, of which the wail of the cell 
consists. 

Cenchium. A name of the peculiar four- 
parted fruit (or the four nutlets around 
a common style) which distinguishes 
Labiate and Borraginacee. 

Centiifjugal. Tending or developing 
from the centre outward. 

Centripetal. Tending or developing from 
without toward the centre. 

Cephalanthium. Synonym of Antho- 
dium. 

Ceratium. A siliquiform capsule, such 
as that of Corydalis, Cleome, &c. 

Cereal. Belonging to corn and the 
allied grains. 

Cerinus. Of the color of wax. 

Cérnuous (-uus). Nodding. 

Chet. Greek for a bristle, Latin Seta. 

Chaff. Small scales; dry and depau- 
perate bracts; such as those on the re- 
ceptacle of asunflower and many other 
Compositze ; also glumes of Grasses. 

Chaffy. Provided with or having the 
texture of chaff. 

Chalaza. The part of an ovule where 
coats and nucleus are confluent; 277. 
Channelled. Hollowed out longitudi- 

nally like a gutter. See Canaliculate. 

Character. A diagnostic description, 
or the enumeration of essential differ- 
ences; 361. 

Chasmoyamy. The opening of the peri- 
anth at flowering time; the opposite 
of Cleistogamy 

Chartaceous (-eus) 
of writing-paper. 

Chlorophyll. The green matter of leaves 
and other vegetation ; 76, 88. 

Chloros. Greek for green. Enters into 
compounds, such as Chlorunthus, 
green-flowered, Chloranthy, same as 
Chlorosis, as when petals turn green; 
12. 

Chlordsis. Literally becoming green, as 
some flowers in retrograde metamor- 
phosis. Also used contrariwise for 
the loss of a normal green color; 172. 

Chorda Pistillaris. A line of tissue reach- 
ing from stigma to ovary. 

Choripetalous (-us). Same as Polypeta- 
lous, 7. e. petals unconnected : 244. 

Chorisepalous. Same as Polysepalous.&e. 

Chorisis. The separation of a leaf or 
phyllum into more than one ; 202. 

Cher’stophyllus. Separate-leaved. 


Having the texture 


GLOSSARY. 


Chromule (-ulz). Coloring matter of 
plants other than chlorophyll, espe- 
cially that of petals. 


Chrysus. Greek for golden, or golden- 
yellow; as 
Chrysanthus. Yellow-flowered, &c. 


Cicatrixz, Cicatricula. A scar left by 
the fall of a leaf or other organ. 

Ciliate (-atus), Ciliaris. Marginally 
fringed with hairs. 

Cilium, pl. cilia. Marginal hairs, form- 
ing a fringe, like the eyelash. (The 
name has been extended in scientific 
books to undividual hairs, and of a 
surface as well as edge.) 

Cincinnus. A’curl: name of a uniparous 
scorpioid cyme, which is Cineinnal ; 
156, 157. 

Cinenchyma. Laticiferous tissue. 

Cinerascens, Cincraceus. Ash-grayish. 

Cimereous (-eus). Ash-gray. 

Cinnabarinus. Cinuabar-color ; searlet 
touched with orange ‘ 

Circinal (-alis). Involute from the tip 
into a coil; 133. 

Circinate, or Circinnate (-atus). Same 
as preceding; or sometimes meaning 
coiled into a ring only. 

Circumscissile, or Circumcissile ( Cireum- 
scissus). Cut cireulary and transverse- 
ly; divided transversely ; 231, 293. 

Circumscription (-iv). The general out- 
line of the margin of a flat body. 

Cirrhiferous (-us) and Cirrhose. 
dril-bearing. 

Cirrhus. A tendril; 54. 

Citreus, Citrinus. V.emon-colored. 

Clados. Greek for branch; whence such 
terms as 

Cladodium. Same as Cladophyllum. 

Cladophyll, Cladophylla. Branches as- 
suming the form and function of foli- 
age: 65, 66. 


Ten- 


Claivate (-atus), Claviformis.  Club- 
shaped. 
Clivellate. Diminutive of Clavate. 


Claviculate (-atus). Furnished with clar- 
icule ; viz, tendrils, hooks, or other 
appliances for climbing. 

Class, 325. 

Classification, 315. 

Clathrate (-atus). Latticed. 

Claw. The narrowed base or stalk which 
some petals, &c., possess; 245. 

Cleistégamy, Cleistogamous, Cleistogamic. 
Close-fertilization in unopened blos- 
soms ; 241. 

Cleistégeny, Cleistogenous, 241. Same as 
Cleistogamy. 


GLOSSARY. 


Cleft Cut half-way down or there- 
about; ¥8. 

Climbmy. Rismg by laying hold of 
surruunding objects for support; 51. 
Clinandrium. ‘The anther-bed in Orchi- 

dacex. 

Clinanthium. A name for the receptacle 
of inflorescence in Composite; 148. 
Clinium. Used in Greek compounds for 
receptacle, e.g. Periclinium, for an 
involucre around the receptacle of 

inflorescence. 

Close-fertilization. Fecundation by own 
pollen ; 216, 280. 

Cloves. A gardener’s name for young 
bulbs developed around a mother bulb. 

Club-shaped. Gradually thickened up- 
ward from a slender base. 

Clustered. Collected in a bunch of any 
sort. Cluster is a good indefinite name 
fur any assemblage of flowers on a 
plant. 

Clypeate (-atus), Clypeiformis. 
ler-shaped. 

Coucerrate (-atus). 


Buck- 


Heaped together. 


Coddnate (-atus), Coadunatus. Same as 
Adnate. 

Coalescence. Union of similar parts; 
179, 180. 


Coalescent (-ens), Coalitus. Cohering; 
properly applied to the organic cohe- 
sion of similar parts. 

Coarctate (-atus). Crowded together. 

Coated. Composed of layers as an 
onion, or furnished with a covering 
or rind. 

Cobwebly. Bearing long and soft entan- 
gled hairs. 

Coccineus. Bright red or scarlet (red 
with a little yellow). 

Coccus. Greek for a kernel or nutlet, 
from which the Latin Coccum, the 
kermes or scarlet grain (supposed 
berry) of the Quercus coccifera; used 
botanically, mostly in the form of 
“coccus,”’ for the portions into which 
a schizocarp, or lobed fruit with one- 
seeded cells, splits up; these portions 
are Cocci or Coccules; 296. 

Cochlear (Cochlearis). Spoon-shaped. 
Unmeaningly applied also to a form 
of imbricative sxstivation with one 
piece exterior; 137. 

Cochleate (-atus). Shell-shaped, i. e. 
spiral in the manner of a snail-shell. 
Celospéermous (-us). Hollow-seeded; ap- 
plied to seed-like carpels of Umbelli- 
ferze with ventral face incurved at top 

aud bottom, as in Coriander. 


403 


Cenanthium. Synonym of Clinanthium. 

Cenobio. Synonyin of Carcerulus. 

Cohesion. ‘Vhe congenital union of one 
organ with another; either similar 
parts (coalescence), or dissimilar parts 
(adnation ). 

Cohort. In classification a group next 
superior to order, 326. 

Coleorhiza. Root-sheath; the invest- 
ment (belonging to the cotyledon or 
plumule) through which the primary 
root in many Monocotyledons bursts 
in germination; 26. 

Collar (Cullum). Name of an imaginary 
something intermediate between pri- 
mary stem and root. 

Collateral. Standing side by side. 

Collective Fruits. The aggregation of 
the fruits of several flowers into one 
mass; 301. 

Colored. Of other color than the green 
of herbage; 118. 

Columella. The persistent axis of cer- 
tain capsules, spore-cases, &c.; 289. 
Column (Columna). Body formed by 
the union of the filaments among them- 
selves (as in a malvaceous flower), or 
with the style or stigma, as in Or- 

chids; 290. 


Columnar. Column-shaped ; _ pillar- 
shaped. 
Coma. Literally a head of hair; a tuft 


of hairs of any sort ; specially a tuft of 
hairs on a seed: 306 Also the name 
of the whole head of a tree. 

Commissure (-ura). The face by which 
two carpels cohere, as in Umbellifere. 

Common ( Communis). General or prin- 
cipal, as opposed to partial. 

Coumose (-osus), sometimes 
Furnished with a coma. 

Cémplanate (-atus). Flattened. 

Complete (Completus). Having all the 
parts belong to it or to the type; 175. 

Complicate (-atus). Folded upon itself. 

Compound. Said of similar parts aggre- 
gated into a common whole. Com- 
pound Flower, 147. Compound Pistil, 
263. Compound Inflorescence, 159. 

Compound Leaf. One divided into sep- 
arate blades; 100. 

Compressed (-us). Flattened lengthwise. 

Concaulescence. A name for the coales- 
cence of axes: 158. 

Conceptacle (-aculum). Originally used 
by Linneus for what is now called 
Follicle; and later for the pair of fol- 
licles of Asclepiadacee and Apocy- 
nacez. 


Comatus. 


404 


Conchiformis. Shaped like one valve of 
a bivalve shell. 

Concinnus. Neat or elegant. 

Concolor. Of the same or of uniform 
color. 

Conduplicate (-atus), Conduplicativus. 
Volded together lengthwise; 133. 

Cone. See Strobile. 

Conjertus. Closely packed or crowded. 

Conferruminate (-atus). Stuck together 
by adjacent faces, as the cotyledons of 
Horsechestuut; 314. 

Confluent (-ens). Blended into one; pass- 
ing by degrees the one into the other. 

Conformed (-ormis). Similar to in form; 
or closely fitted to, as a seed-coat to 
the nucleus. 

Congested (-us). Crowded together. 

Conglobate (-atus). Collected into a ball. 

Conglomerate (-atus). Densely clus- 
tered or heaped together. 

Coniferous (-us). Cone-bearing. 

Conjugate (-atus). Coupled; in single 
pairs. Conjuyate-pinnate, 104. 

Connate (-atus). United congenitally; 
107, 182. 

Connate-perfoliate. United at base in 
pairs around the supporting axis; 108. 

Connective (-ivum). 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; 238 


Consolidated. When unlike parts are 
coherent. 
Continuous. The reverse of articulated 


or interrupted. 

Contorted (-us). Twisted; or bent or 
twisted on itself. In /Estivation, the 
same as Conyolute; 138. 

Contortuplicate (-atus). 
plaited or folded. 

Contracted. Either narrowed or short- 
ened. 

Contrary (-arius). Opposite in direction 
to the part compared with; asa silicle 
compressed contrary to the dissepi- 
ment. 

Convolute (-utus) or Convolutive (-ivus). 
Rolled up from the sides or longitudi- 
nally. In stivation, 138. In Ver- 
nation, 133. 

Céralloid (-eus). Coral-like. 

Céreulum. Old name for the embryo, 
or Cor seminis; 311. 


Twisted and 


GLOSSARY. 


Cordate (-atus), sometimes Cordiform 
(-ormis). Heart-shaped; like the fig- 
ure of a heart on cards; the stalk at 
the broader and notched end ; 96. 

Coriaceous (-eus). Leathery in consist- 
ence. 

Cork, 81. 

Corky. Of the texture of cork. 

Corky Envelope, 76. 

Corm (Cormus). A  bulb-like fleshy 
stem, or base of a stem; a “solid 
bulb;” 61. 

Cormophytes ( Cormophyta), 341. 

Corneous(-eus). Of the texture of horn. 

Corniculate (-atus). Furnished with a 
little horn. 

Cornu. A horn; i.«. a horn-like process; 
sometiynes used for Calcar, a spur. 

Cornute (-utus). Furnished with a horn- 
like process or spur. 

Corolla. The interior perianth, com- 
posed of petals; 165, 243. 
Corollaceous (-eus), Corollinus. 
ing to, or resembling corolla. 
Corolliferous (-us). Bearing a corolla. 

Corclliflorous, Corolliflore, 340. 

Coréllula. Dimiuutive of corolla. 

Corona. A crown: an inner appendage 
to a petal, or to the throat of a corolla; 
210, 246. Orany coronet-like append- 
age at the summit of (crowning) an 
organ. 

Céronate (-atus). 
corona, &e. 

Coréniform (-ormis). 
crown or coronet. 

Cérrugate (-atus or -ativus). 
or in folds; 133. 

Cortex. Rind or bark. 

Cortical (-alixs). Relating to bark. 

Corticate (-atus). Coated with a bark 
or with an accessory bark-like cover- 
ing. 

Cérymb (Corymbus). A flat-topped or 
merely convex and open flower-cluster 
of the indeterminate or centripetal 
order; 146. 

Corymbiferous (-us). Bearing corymbs. 

Cérymbose. In corymbs, or in the man- 
ner of a corymb. The corymb of 
Linneus and of other writers down 
to Roeper included most cymes. So 
that much cymose inflorescence is 
in descriptions loosely said to be 
corymbose, or a stem is said to be 
corymbosely branched, even when 
the evolution is centrifugal ; 146. 

Costa. _A rib; when single, a midrib 
or mid-nerve. ae 


Pertain- 


Crowned, having a 
Shaped like a 


Wrinkled 


GLOSSARY. 


Costal-nerved. With nerves springing 
from a midrib; 92. 

Costate (-atus). Ribbed; furnished with 
one or more lorgitudinal primary 
veins or ribs. 

Cotylédons ( Cotyl’don, pl. 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. : 

Cratéviform (-ormis). In the shape of 
a goblet or cup, of hemispherical con- 
tour or more shallow; 248. 

Cr*mocarp (Cremocarpium). A dry and 
seed-like fruit, composed of two one- 
seeded carpels, invested by an epigy- 
nous calyx, and separating at matu- 
tity; 297. 

Creeping. Running along or under 
ground and rooting; 53. 

Crena, Crenatura. A rounded tooth or 
notch. 

Crenate (-atus). Toothed by crenatures : 
scalloped; 98 

Crenei, Crenelled. Same as Crenature 
and Crenate. 

Crénulate (-atus). Diminutive of Cre- 
nate, 2. ¢. with small crenatures. 

Crested. Yurnished 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. 

Crispatus. Curled or crispy. 

Cristate (-atus). Crested. 

Croceus, Crocatus. Safiron-colored, 7. e. 
deep reddish-yellow. 

Cross-breeds. The progeny 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). 
suinmit of an organ. 

Cruciate (-atus), Cruciform (-ormis). 
Cross-shaped. 

Cruciferous (-us). Cross-bearing: used 
in the sense of Cruciform; as the 


Borne on the 


1 


405 

“‘cruciferous’’ corolla of the order 
Crucifere ; 246. 

Crumpled. See Corrugate. 

Crustaceous (-us)., Of hard and brittle 
texture. 

Cryptos. Greek for concealed; whence 

Cryptogamia. Cryptogamous or Cryp- 
togamic plants ; 3, 335, 544. 

Cryptégamous. Pertaining to the above. 

Cucullate (-dtus), Cucullaris, Cuculli- 
Jormis. Hooded, or hood-shaped, 
cowled. 

Culm (Culmus). The peculiar stem or 
straw of Grain-plants and Grasses; 40. 

Cultrate (-atus), Cultriformis. 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 iike; 296. 
Cupularis, Cupulatus. Furnished with 
or subtended by a cupule or any re- 

sembling body. 

Cupuliferous (-us). Cupule-bearing. 
Curvinerved (-ius). When the ribs of a 
leaf are curved in their course; 92. 
Curviserial. 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). 

or pellicle. 

Cutting. A severed portion of a plant 
used for bud-propagation: 43. 

Cydneus. A clear bright blue. 

Cyathiform (-ormis). Cup-shaped; in the 
form of a 

Cyathus. A drinking-cup, such as a 
goblet or wine-glass. 

Cycle. A circle. Sometimes used for 
one turn of a helix or spire; 122. 

Cyclical. Relating to acycle; or coiled 
into a cirele; 119, 120. 

Cylindraceous. Somewhat or nearly 
cylindrical. 

Cyl: ndrical (-us). Elongated and with 
circular cross-section; in the form of a 
cylinder. 

Cymbeform or 
Boat-shaped. 

Cyme ( Cyma). 
determinate 


The acorn-cup and 


The outermost skin 


Cymbiform (-ormis). 


A flower-cluster of the 
or centrifugal type, 


406 


especially a broad and flattish one; 
151. 

Cymo-bétryose. When cymes are ar- 
ranged in botryose manner; 159. 

Cymose (-osus). Bearing: cymes, or re- 
lating to a cyme; 181. 

Cymule (Cymula). Diminutive ecyme, 
or a portion of a cyme; 151. 

Cynarrhodium. Name of such a fruit as 
that of the Rose; fleshy, hollow, and 
euclosing achenia. 

Cypsela. Name of an ache: ium in- 
vested by an adnate calyx, as the fruit 
of Composite; 295- 

Cystclith. Onz of the mineral and 
usually partly crystalline concretions 
of the cells of the epidermis of or 
subjacent tissue of the leat in various 
plants, especialiy in Urticacez. 

Cytoblast. An obsolete name for the 
nucleus of a cell of cellular tissue. 


Dactylose (-osus). Fingered, or finger- 
shaped. 

Dasyphyllous (-us). Woolly-leaved. 

Deald ite (-atus). Whitened over (as if 
whitewashed) with a white powder or 
minute pubescence. 

Deca. Greek for ten, compounded with 
various words, such as 

Deengynia. One of the Linnean artificial 
orders; 337. 

Decayynous (-us). 
carpels. 

Decamerous(-us). Of ten members; 176. 

Decandria. A Linnean class with ten 
stamens: 334. 

Decandrous (-us). 
249. 

Decapétalous (-us), Decasepalous, &c. 
With ten petals or sepals, &c. 

Deciduous (-us). Falling, or subject to 
fall in season, as petals after anthesis, 
and leaves (except of evergreens) in 
autumn; 245. 

Déclinate (-atus), or Declined. Bent or 
curved downward or forward. 

Decompound. Several times compound- 
ed or divided; 102, 104. 

Decompositus. Decompound. 

Decumbent (-ens). Reclining, but with 
summit ascending; 53. 

Decurrent (-ens), Decursive. Running 
down into; as where leaves are seem- 
ingly prolonged below their insertion, 
and so run down the stem. 

Decussate (-atus). In pairs alternately 
crossing at right angles. 


With ten styles or 


With ten stamens; 


GLOSSARY. 


Deduplication, Fr. Dédoublement. Same 
as Chorisis; 202. 

Definite (-itus). Of a fixed number, not 
exceeding twenty ; or of a fixed or. er. 

Definite Inflorescence. Where axes of 
inflorescence end ina flower; 144, 151. 

Dejflexed (-us). Bent or turned abruptly 
downward. 

Dejflorate (-atus). 
state. 

Defoliate (-atus). Having cast its leaves. 

Defoliution, 87. : 

Dehiscence (-entia). The mode of open- 
ing of a capsule or anther by valves, 
slits, or regular lines; 288. 

Dehiscent (-ens). Opening by regular 
dehiscence; 292. 

Deliquescent (-ens). Dissolving or melt- 
ing away, as a stem divided into 
branches; 48. 

Deltoid (-oides). Having the shape of 
the Greek letter A. 

Demersed (-us). Under water; same as 
submersed. 

Dendritic (-icus), Dendroid (-oideus). 
Tree-like. 

Dendron. 


Past the flowering 


Greek for tree. 

Deni. Ten together. 

Dens. A tooth. 

Dentate (-atus). Toothed; specially-with 
salient teeth not turned forward ; 98. 

Denticulate (-atus). Minutely toothed ; 
having denticulations, or diminutive 
teeth. 

Denidate (-atus). Made naked; stripped. 

Deorsum. Downwards. 

Depauperate (-atus). Tmpoverished; as 
if starved; or diminutive for want of 
favorable surroundings. 

Depressed (-us). Having the appear- 
ance or shape as if flattened from 
above. 

Derma. Greek for skin or surface of a 
plant or organ. 

Descending (-ens). Tending or turning 
gradually downward. 

Descending Axis. Primary root; 11. 

Determinate. Limited in number or ex- 
tent; as are the axes of determinate 
inflorescence; 144, 151. 

Désinens. Terminating in. 

Desmus. Greek for things bound, or as if 
chained together. 

Dextrorse (Dextrorsus: adv. Deztror- 
sum). Toward the right hand, or re- 
lating to it; 51, 140. 

Di, Dis. In Greek compounds, two, or 
double. 

Diachenium. Synonym of Cremoearp. 


GLOSSARY. 


Diadelphia. A Linnean (335) class 
having the stamens. 

Diadelphous (-us). Combined by their 

_ filaments into two sets; 250. 

Diagnosis. A brief distinguishing char- 
acter. 

Dialypetale, 341. 

Dialypetalous (-us). Same as poly peta- 
lous, t. €. of separate petals; 244. 

Dialyphyllous (-us). Bearing separate 
leaves. 

Diandria. A Linnean class with per- 
fect flowers having only two stamens; 
334. si 

Diandrous ( Diander, &e.). Having two 
stamens; 249. 

Diaphanous (-us). 
shine through. 

Dicarpellary. Composed of two carpels 
or pistil-leaves; 261 

Dichasium. A two-parted or two-rayed 
eyme; 152, 155. 

Dichlamydeous (-eus). Having a double 
perianth; 191. 
Dichétomous (-us). 

two-forked. 

Dichéyamous (-us), Dichogamy. Her- 
maphrodite with one sex earlier de- 
veloped than the other in the blossom; 
219. 

Dicksium. Name of a fruit consisting 
of an achenium within a separate and 
free covering made of perianth, as that 
of Mirabilis. 

Diclinvus (Diclinis). When flowers are 
of separate sexes; 191. 

Dicéccous (-us). Fruits of two cocci. 

Dicotyl-dons, Dicotyledones. Plants of 
the class marked by having two coty- 
ledons; 27, 339, 340, 544. 

Dicctyledonous (-eus). Having a pair of 
cotyledons; 10, 314. 

Didymous (-us). Twin, found in pairs. 

Didyndmia. The Linnean class marked 
by didunamy (335), 1. e. 

Didynamous (-us). When a 4-androus 
flower has the stamens in two pairs, 
aud one pair shorter than the other; 
250. 

Dier¢silis. 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. 
Digamous (-us). 
same cluster. 


Letting the light 


Forked in pairs; 


Of two sexes in the 


407 


Digitate (-atus). Fingered; a compound 
leaf in which all the leaflets are borne 
on the apex of the petiole; 101. 

Digitately. In a digitate mode; same 
as Palmately. 

Digitate-Pinnate, 104. 

Digynia. A Linnzean order character- 
ized by having the gyneecium 

Digynous. With two separate styles or 
carpels; 261. 

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. 
Diecia. Linnean class (355) of plants 


with the flowers 

Diecicus (Dizcius, Dioicous). Unisex- 
ual, and the two sexes borne by dis- 
tinct individuals; 191. 

Diecio-polygamous. When some indi- 
viduals bear unisexual and others bi- 
sexual flowers. 

Dipetalous (-us). Two-petaled; 244. 

Diphyllous (-us). Two-leaved; 243. 

Diplo. See Duplo. 

Diplostémonous, Diplostémeny. Having 
twice as many stamens as petals or 
sepals; 177, 198. 

Diplotegium. <A capsule or other dry 
fruit, invested with adnate calyx; an 
inferior capsule. 

Dipterous -(us). Two-winged. 

Diremption (-io). Syn. of Chorisis; 202. 

Disciferous (-us). Disk-bearing. 

Disciform (-ormis). Depressed and cir- 
cular, like a disk or quoit. 

Discoidal or Discoid ( Discoideus). Ap- 
pertaining toa disk. A discoid head 
is one destitute of ray-flowers. 

Disc or Disk (Discus). A word used in 
several senses. The disk or dise of a 
flower is a development of the torus 
within the calyx, or within the corolla 
and stamens; 213. In acapitulum 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- 
cent. 


408 


Disepalous (-us). Two-sepaled; 244. 

Disk-jlowers. Those belonging to the 
disk, or body, and not to the margin 
or ray of a capitulum. 

Dissected (-us). Deeply cut or divided 
into numerous segments. 

Dissépiment (-éntum). A partition in an 
ovary or pericarp; 264. 

Dissilient (-ens). Bursting asunder or 
in pieces. 

Distichous (-us}. Disposed in two ver- 

, ticalranks; 122. 

Distinct ( Distinctus). 
not united. 

Distractile (-ilis). Carried widely apart. 

Diihécous (-us). Of two thecee, or cells, 
as are most anthers; 254. 

Diurnal. Daily ; occurring in the day; 
sometimes used for ephemeral. 

Divaricate (-atus). 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 

Dodecagynia. Linnan order with flow- 
ers. 

Dodecagynous. Having twelve styles or 
distinct carpels. 
Dodecamerous (-us). 

the circle. 

Dodecandria. A Linnean class (334) 
with the flowers. 

Dodecdndrous. Having twelve (or from 
12 to 19) stamens; 249. 

Dodrantalis. A span (about nine inches) 
long. 

Dolubriform (-ormis). 
hatchet-shaped. 

Dorsal (-alis). Relating to the dorsum 
or back. 

Dorsal Suture. That which answers to 
the midrib of a carpel; 261. 

Dorsiferous. 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 flower are monstrously increased 
mostly at the expense of the essential 
organs. 

Downy. Pubescent with fine nd soft 
hairs. Loosely synonymous with soft- 
pubescent, tomentose, Ke. 

Drepanium. A sickle-shaped eyme; 156. 

Drupaceous (-eus). Resembling or relat- 
ing to a drupe. 

Drupe (Drupa). A stone fruit; 297. 

Drupelet, Drupel (Drupéola). A dimin- 
utive drupe; 297. 


Separate from; 


Of twelve parts in 


Axe-shaped or 


GLOSSARY. 


Drupetum. An aggregation of drupes; 
300. 

Duct. In vegetable anatomy, an elon- 
gated cell or tubular vessel, ound espe- 
cially in the woody (fibro-vascular) 
parts of plants. 

Dimetose (-osus). Pertaining to Dume- 
tum, a thicket, or Dumus, a bush. 

Dumose (-osus). Bushy, or relating to 
bushes. 

Duplo. Twice as many. In Greek com- 
pounds, Diplo. 

Duramen. ‘The heart-wood of an exo- 
genous stem ; 80. 

Dwarf. Of small size or height com- 
pared with its relatives. 

Dyclesium. See Diclesium. 


E-or Ezx- As a prefix to Latin words, 
carries a privative meaning, as Ecos- 
tate, without ribs, Exalbuminous, with- 
out albumen. 

Eared. Same as Auriculate. 

Ebracteate, Ebracteolate (-atus). 
tute of bracts or bractlets. 

Eburneus. Ivory-white. 

Ecalcarate (-atus). Spurless. 

Echinate (-atus). Beset with prickles, 
like a hedgehog. 

Echinulate (-atus). 
tive prickles. 

Edentate (-atus). Toothless. 

Lffete (-etus), or Effetus. Past bearing; 
functionless from age. 

Efflorescence (-entia). The time or state 
of blossoming; anthesis. 

Effuse (-usus). Very loosely spreading, 
more so than diffuse. 

Eylandulose (-osus). Destitute of glands. 

Eqgy-shaped. See Ovate. 

Eladter. (ne of the spiral or spirally- 
marked threads in the spore-cases of 
certain Hepatice. 

Eldtus. Tall or lofty. 

Eleutheros. In Greek compounds, sep- 
arate or distinct. 

Eleutheropetalous (-us). Same as Chori- 
petalous or Polypetalous; 245. 

Ellipsoidal (-eus). An elliptical solid; 
sometimes used for nearly elliptical. 

Elliptical (-us). In the form of an el- 
lipse. Oval or oblong with regularly 
rounded ends: 95. 

Emarcidus. Flaccid or withered. 

Emarginate (-atus). With a notch cut 
out of the margin; or, as usually ap- 
plied, out of the extremity; 97. 

Embracing. Clasping by the base. 


Desti- 


Beset with diminu- 


GLOSSARY. 


Embryo or Embryon. The rudimentary 
plautlet formed in a seed; 9, 311. 

Embiyonal. Relating to the embryo; 
as Lmbryonal Vesicle ; 28+. 

Embryo-sac. The cell in the ovule in 
which the embryo is formed; 283. 

Embryogeny. Embryo-formation. 

Emersed (Emersus). Raised above and 
out of the water. 

Enantioblastus. With embryo at the end 
of the seed diametrically opposite the 
hilum. 

£nation. Having outgrowths from the 
surface, &e.; 179. 


Endeca. In Greek compounds, eleven; 
as in 
Endecandrous, Endecayynous. With 


eleven stamens or eleven styles, Ke. 
Endemic. Contined geographically to 
the particular region. 
Endocarp (-drpium). The inner layer of 
_a pericarp; 288 
Endechrome (-oma). Peculiar coloring 
matter in cells; especially the color- 
, ing matter of Algw. 
Endogens, Lndogene. 
Plants; 70. 
Endogenous structure, 70. 
Endopleura. Inner seed-coat; 306. 
Endophleum. 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 
Endorhize. 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. 
or veins visible. 


Endogenous 


Nerveless; no ribs 


Ennea. In Greek compounds, nine; 
as in 

Enneagynia. A Linnean ordinal name, 

> and 


Ennedgynous. With nine separate styles 
or carpels; 337. 

Enneandria. Linnean class, and Enne- 
androus, with nine stamens; 249, 334. 

Enédal (Enodis). Without a node. 

Fnsatus. Same as Ensiform. 

Ensiform (-ormis). Sword-shaped ; 7. e. 
like a_broad sword, or the leaf of an 
Iris. es? 


409 


Entire. Without toothing or division; 
the margin whole and even; 97. 

Entomophilous. said of flowers which 
are habitually fecundated by pollen 

, carried by insects ; 217, 218. 

Entophytes (Entophyta). Plants grow- 
ing in or out of other plants, as cer- 
tain Fungi, &e.; 4. 

Ephemeral. Lasting only for one day. 

£pi. In Greek compounds, upon. 

Lpiblast (-us). Name sometimes given 
to the first (and an undeveloping) leaf 
of the plumule of the embryo of grasses 
and grain. 

Lpiblastéma. A superficial outgrowth 
from leaves, &c.; 210. 

Epicalyz. Name sometimes given to 
an involucel resembling an accessory 

_ calyx. 

Epicurp (Epicarpium). The external 
layer of a pericarp; 288. 

Epichilium. The terminal portion of 
the labellum of an Orchid, when this 
is of two parts. 

Epiclinal (-us). Upon a torus. 

Epicérolline. Upon a corolla. 

Epidermis. The skin of a plant; 76, 89. 

Epigeous (-eus) Growing on or out of 
the ground. 

Epigynous (-us). Literally on the pistil; 
meaning on the ovary, or seemingly 

185. 

Epipetalous (-us). Borne on (adnate to) 
the petals; also used in the sense of 
piaced before the petals. 

Epiphizum. The outermost or corky 
bark; 76. 

JF piphyllous (-us).~ Growing on leaves. 

Fpiphytal. 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. 

Episepalous. On the sepals; also used 
in the sense of standing before a 

_ sepal. 

Episperm ( lpispermium). 

_ outer coat of a seed; 305 

Epitropors (-us). Name (by Agardh) of 
an aratropous ovule with rhaphe 
averse when ascending, adverse when 
suspended: 282. 

Equal (qualis). Alike as to length or 

,number, &c., as the case may be. 

Equitant '-ans). Riding: folded around, 
as if straddling over; 108, 138: 

Erect (£rectus). Standing upright, 
mostly in relation to the ground, 


sO; 


The coat or 


410 


sometimes when perpendicular to the 
surface of attachment. 

Erion. Greek for wool; used in com- 
pound words, as 

Evianthus. With woolly flowers. 


Erioéphorus. Wool-bearing. 
Eriophyllus. Woolly-leaved. 
Erostrate (Erostris). Beakless. 


Erose (Lrosus). As if gnawed; applied 
to an irregularly toothed or eroded 
margin. 

Erythros. Greek for red, used in com- 
pound terms from the Greek. 

Erythrostomum. Name given by Des- 
vaux for such an aggregate fruit as 
a raspberry; 300. 


Estivation. See #stivation. 
cterio. Name of aggregate fruits, 


especially of fleshy ones, such as a 
blackberry. 

Etivlated. Blanched by darkness. 

Eu. Prefixed to words of Greek deri- 
vation denotes very, true, or much so. 
Frequently used in names of sections 
or other groups; 357. 

Euphylia. Foliage-leaves, or true leaves. 

Lutropic (-icus). Name suggested for 
twining ‘‘with the sun;’’ dl. 

Evalvular (Evalvis). Not opening by 
valves. 

Evergreen. Bearing green foliage all 
the year rcund. 

Evittatus. Not vittate. 

Ex. A prefix in place of £ privative 
when the following part of the com- 
pound begins with a vowel; as 

Exalbuminous (-osus). Destitute of albu- 
men; 14, 309. 

Exalate (-atus). Destitute of wing. 
Ezxanthemata. Eruptive excrescenses on 
the surface of leaves, &e.; blotches. 
Exaristate (-atus). Destitute of an 
arista or awn. 
Exdsperate (-atus). 

projecting points. 

Excentric (-icus). 
one-sided. 

Excurrent (-ens). Running through to 
the very summit or beyond; 48. 

Exiguus. Small or mean. 

Ezilis. Lank or meagre. 

£zimivs. Distinguished, as for size or 
beauty. 

Fzo-. In Greek compounds, external or 

, outward; as 

Exocarp (Exocarpium). The outer layer 
of a pericarp; 288. 

Exé,enous. Outside growing. as the 
wood of Dicotvledons; 69, 73. 


Rough with hard 


Out of the centre; 


GLOSSARY. 


Exogens, Exogene. Exogenous Plants; 
69, 340. 

Exorhize. Name equivalent to Exogen, 
from 

Ex hizal(-us). The radicle not sheathed, 
so the primary root in germiration 

, has no covering to break through. 

Exostome (Exostoma). The foramen of 
the outer coat of the ovule; 277. 

Exothécium. The outer wall of an 

, anther. 

Explunate ( atus). Spread out flat. 

Exsert, Exserted (Exsértus). Protruding 
beyond or out of, as stamens beyond 

_ the corolla. 

Exstipulate (-atus). Destitute of stipules. 

Exterior. External in the sense of outer. 
But also in the flower sometimes used 
in the sense of anterior. 

FExtine. Outer coat of a pollen-grain. 

Extra-azillary. Beyond or out of the axil. 

Extrorse (Ezxtrorsus, Extrorsum). Di- 
rected outward: 253. 

Eye. A gardener’s name for an unde- 
veloped bud. 


Facies. Face; the general aspect. 

Falcate (-atus), and Falciform (-ormis). 
Scythe-shaped or sickle-shaped; plain 
and curved, with the edges parallel. 

Family. In botany, synonymous with 
Order; 325. 

Fan-shaped. See Flabelliform. 

Farina. Starch. 

Farinaceous (-eus). Of the nature of 
starch, or containing it. 

Farinose (-osus). Covered with a meal- 
like powder. 

Fasciate (-atus). Said of monstrous ex- 
pansions of stems, giving the appear- 
ance as of several stems coalescent in 
one plane. 

Fascicle (-icula). A close cluster or 
bundle, whether of flowers, stalks, 
roots, or leaves; 147, 153. 

Fascicled (Fasciculatus, Fascicularis). 
In a fascicle; 131. 

Fastigiate (-atus). Said of branches 
when parailel, clustered, and erect. 
Faux, pl. fauces. The gorge or throat 
of a gamophyllous calyx or corolla; 
either at the orifice, or a portion 
between the limb and the proper 

tube; 246. 

Faveolate (-atus), Favose (-osus). Honey- 
combed; same as Alveolate. 

Feather-veined. Waving veins all pro- 
ceeding from the sides of a midrib. 

Feathery. See Plumose. 


GLOSSARY. 


Fécula or Fecula. Starch-like matter. 
Applied to a pistillate flower, or to a 
pl nt producing only such flowers. 

Fenistrate (-atus), Fenestralis. Pierced 
with large holes, like windows. 

Ferrugineous or Ferruginous (Ferru- 
gineus). Colored to imitate iron-rust. 

Fertile (-ilis). Fruitful, fruiting, or ca- 
pable of producing fruit; as a fertile 
flower is one provided with a well- 
formed pistil; 191. In English deserip- 
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 (Fibra). Any fine filament; the 
elementary components of wood, &c. ; 
delicate roots, Ke. 

Fibril (-illa) A diminutive fibre. 

Fibrillate (-atus), Fibrillose  (-osus). 
Furnished or abounding with fibres 
or fibrils. 

Fibrous, Fibvose (-osus). 
of the nature of fibres. 

Fibro-vasculur. 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 ananther; 165,251. Also any 
fibre-shaped or thread-like body. 

Filamentous, Filamentose (-osus). 
posed of threads or filaments. 

Filicology. The botany of Ferns: re- 
placed by Pteridology. 

Filiform (-ormis) . Thread-shaped; long, 
slender, and terete. 

Filipéndulous (-us). 
thread, 

Fimbria. A fringe, or dissected border. 

Fimbriate (-atus). Fringed ; bordered 
by slender processes or marginal ap- 
pendages 

Fimbrillate (-atus), Fimbriliferous (-us), 
Bearing Fimbrille 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, Fistulose (-osus). Hollow 
through the whole length, as the leaf 
and stem of an Onion. 

Fldbellate (-atus), Flabelliform (-ormis). 
Fan-shaped; much dilated from a 


Composed or 


Com- 


Hanging from a 


| 


411 


wedge-shaped base, and the broader 
end rounded, 

Flabellinerved. With radiating straight 
nerves; 92. 

Flaégellute (-atus), Flagellaris, Produe~ 
ing filiform runners (flagella), or 
runner-like branches. 

Flagelliform  (-ormis). Runner-like; 
long, slender, and supple like a whip- 
lash or Flagellum ; 53. 

Flammeus. \lame-colored. 

Flavescent (-ens). Yellowish or pale 
yellow. 

Flavus. Pale yellow or ochre-yellow. 

Fleshy. Succulent; of the consistence 
of flesh. 

Flexuous, Flexuose (-vsus). Zigzag; 
bent alternately im opposite direc- 
tions. 

Floating. Borne on the surface of water. 

Floccove (-osus). Bearing or clothed with 
locks of soft hairs or wool ( flocc7). 

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. 

Floral. Belonging to the flower. 

Floral Envelopes. YFlower-leaves; 164. 

Floret. A small flower, one of a cluster. 

Flovibundus. Abundantly floriferous. 

Flovifcrous( Florifer, Floriferus). Bear- 
ing flowers 

Flérula. A small Flora; the Flora of 
a restricted district. 

Flos. Latin for flower. — Flos plenus. A 
“ double’? flower; that is, one in which 
petals are increased abnormally, com- 
monly at the expense of the andree- 
cium or the gyneecium also; 171. 

Floésculus. Latin for floret. 

Flower. The whole reproductive appa- 
ratus in a phenogamous plant; 163. 
Flower-bud. An unexpanded blossom 

or undeveloped cluster; 40. 

Flowering Plans, 3, 344. 

Flowerless Plints, 3, 344. 

Fluitans. Floating. 

Fliviatile, Fluviatilis. Belonging to a 
river or running water. 

Fly-traps, 113 

Femineus. Feminine or female flower, 
plant, &e.; 191. 

Foliaceous (-eus). Leaf-like in texture 
or appearance; or bearing leaves. 

Foliar (Foliaris). Relating to leaves. 

Foliation (Feliatio). WLeafing out. 

Foliate (-atus). Waving leaves. With 
Latin numerical prefix, 5/foliale, tri- 


412 


foliate, and so on, according to the 
number. 

Foliijvrm (-ormis). Leaf-shaped. 

Foliolate (-atus). Having leaflets: their 
number may be indicated by Latin 
numerals, as b/foliolate, trifoliolate, 
&e.; 102. 

Foliilum. <A Jeaflet; 102. 


Folivse. Bearing numerous leaves. 
Folium. Latin tor leaf; 85. 


Follicetum. A whorl or aggregation of 
follicles; 300. 

Follicle (-iculus). Fruit of asingle carpel 
dehiscent by one (the ventral) suture; 
292. Name of the earlier botanists 
for any kind of capsular fruit. 

Follicular (-aris). Pertaining to or like 

- a follicle. 

Foot-stalk. Petivle, 85; or Peduncle, 143. 

Fordmen. An aperture of any kind; 
specially that of the coat of the ovule; 
277. 

Foraminulose (-osus). Pierced with many 
small holes. 
Forcipate (-atus). 

cers. 

Forked. Divided equally into branches. 

Fornicate (-atus). Arched over, as by 
scales (Fornices) covering the throat 
of the corolla of Hound’s-tongue, &e. 

Foveate (-atus), and diminutive Foveo- 
late. Pitted; impressed with shallow 
depressions or pits, Fovee. 

Foville. Minute granules in a liquid, 
in the protoplasm of the pollen-grain, 
&e.; 258. 

Free. Not adnate to other organs. 
Some imes used in the sense of dis- 
tinct, 7. e. unconnected with others of 
the same sort. 

Fringed. See Fimbriate. 

Frond (Frons). An old name for leaf: 
employed mainly for the leaf of Ferns 
and other Cryptogamia, and certain 
Phenogamous plants which serve for 
fructification as well as foliage; also 
for the peculiar foliage of Palms; 67. 

Frondescence (-entia). The act of leaf- 
ing. Has also been employed to 
express the metamorphosis of floral 
organs into foliage-leaves; 174. 

Frondose (-osus). Sometimes used in 
the sense of leafy; also frond-like, or 
bearing fronds. 

Fructification. The act or the organs 
of fruiting or reproduction through 
flower and seed, or their analogues. 

Fructus. Latin for Fruit. 

Fruit. The immediate product of fruc- 


Like forceps or pin- 


GLOSSARY. 


tification ; in phenogamous plants. 
the seed-vessel and contents, along 
with all intimately connected acces- 
sory parts; 285. 

Fruit-dots in Ferns. See Sorus. 

Frumentaceous. Relating to grain (Fru- 
mentum). 

Frustulose (-osus). Consisting of similar 
pieces or Frustules (frustula). 


Frutex. A shrub. 
Frutescent (-ens). Shrubby, or becom- 
ing so. 


Fruticose (-osus). 
to shrubs. 

Fruticulose (-osus). Relating to a di- 
minutive shrubby plant. 

Fruticulus, A minute or low shrubby 
plant. 

Fugacious. Falling or fading very early; 
lasting a very short time. 

Fulvous (-us). Tawny; orange-yellow 
and gray mixed. 
Fulcra. Accessory organs, such as ten- 
drils, stipules, spines, and the like. 
Fulcrate (-atus). Propped, supported 
by, or provided with accessory organs. 

Fuliginous (-osus). Sooty-brown. 

Fungiform (-ormis and Fungilliformis). 
Mushroom-shaped. 

Fungose (-osus). Spongy in texture; 
fungus-like. 

Finicule, Funiculus. 
ovule or seed; 276. 

Funnelform, Funnel-shaped; 249. 
Infundibuliform. 

Furcate (-atus). Forked; or divergently 
branched. 

Furcellatus. Diminutively forked. 

Furfuraceous (-eus). Scurfy; covered 
with bran-like scales or powder. 

Furrowed. See Sulcate. 

Fuscous (-us). Grayish-brown in hue. 

Fusiform (-ormis). Spindle-shaped ; 
terete and tapering gradually to each 
end; 31. 


Shrubby, or relating 


The stalk of an 


See 


Gdlbulus. The peculiar strobile of Cy- 
press and Juniper, composed of up- 
wardly thickened or fleshy scales; 
393. 

Galea. A helmet; name given, from 
its shape, to the upper sepal of Aconi- 
tum, and the upper lip of certain 
forms of bilabiate corolla; 247. 

Galeate (-atus): Having a galea; hel- 
met-shaped; 247. 

Gumo-. In Greek compounds, denotes 
union by the edges or coalescence. 


GLOSSARY. 


Gamopetalous. A corolla of coalescent 
petals ; formerly Monopetalous ; 244. 

Gamophyllous (-us). Composed of coa- 
lescent leaves. 

Gamosepualous. 
sepals; 244. 

Geitondgamy. Fecundation of a pistil 
by pollen of another flower of the 
same plant; 216. 

Geminate (-atus). 
side by side. 

Gemma. A bud, specially a leaf-bud. 

Gemmation. Budding-growth; or the 
disposition of buds. 

Gemmule (Gémmula). Diminutive of 
gemma; minute and simple buds or 
bodies analogous to buds; also sy- 
nonym of Plumule. Fora time used by 
Endlicher and others for the ovule. 

Genera. Plural of Genus; 323. 

General (-alis). Opposed to partial; as 
general inyolucre. 

Generic, Relating to genus. 

Genetic. Genealogical ; that which comes 
by inheritance. 
Geniculate (-atus). 

a knee. 

Genitalia. The stamens and pistils or 
their analogues. 

Genus. Kind or group superior to spe- 
cies, and which with the species gives 
the name to the plant; 323. 

Geoblast (-astus). A plumule which in 
germination rises from underground, 
such as that of the Pea. 

Germ. A growing point or initial growth, 
as of a bud; or the Embryo; 311. Or 
in the sense of 

Germen. The Linnean name of the 
ovary; 166. : 

Germination (-atio). The act of devel- 
opment of the embryo of a seed into a 
plant. 

Gerontogeous (-eus). 
Old World. 

Gibbous, Gibbose (-osus) Swelling out 
on one side into a gibber or gibber- 
osity. 

Gigantéus. Of unusual height. 

Gilrus.. Dirty yellow with a tinge of red. 

Glabrous ( Glaber). Smooth in the sense 
of not pubescent or hairy. 

Glabrate (-atus). Somewhat glabrous, 
or becoming glabrous. 

Glabriisculus. Almost but not quite 
glabrous. 

Gladiate (-atus). Sword-shaped ; in the 
form of a sword-blade, whether straight 
or somewhat curved. See Ensiform. 


A calyx of coalescent 


Twin; in pairs; two 


Bent abruptly, like 


Belonging to the 


413 


Gland for Glans. An acorn and the 
like; 296. 

Gland (Glandula). A definite secreting 
surface or structure on the surface of 
any part of a plant, or partly imbedded 

" in it, extended to any protuberance 
or structure of similar nature which 
may not secrete. 

Glandular, Glandulose (-osus). Bearing 
glands or having the nature of glands. 

Glanduliferous (-us). Gland-bearing. 

Glareosus. Growing in gravel. 

Glaucescent (-ens). Verging upon or be- 
coming glaucous. 

Glaucous (-us). Covered or whitened 
with a bloom, like that on a Cabbage- 
leaf. 

Globose (-osus). Having or approaching 
a spherical form. 

Globular (-aris), Globulose (-osus). Some- 
what or nearly globose. 

Glochideous, Glochidiate (-atus). When 
bristles and the like are barbed at tip. 

Glochis. A barb. 

Glomerate (-atus). Compactly clustered, 
especially into a 

Glémerule (Glomérulus, Glomus). <A 
cyme condensed into a head or capi- 
tate cluster; 152. 

Glossology, 3, 358. 

Glumaceous (-€us). 
sembling glumes. 

Glume, Gluma. One of the chaff-like 
bracts of the inflorescence of Grasses 
and their relatives; 143. 

Glumella. Diminutive of gluma; an 
inner or secondary glume. 

Glutinous (-osus), Covered with a sticky 
exudation. 

Gonophore (Gonéphorum). A stipe which 
elevates both stamens and pistil; 212. 

Gossypine (-inus). Cottony; flocculent. 

Gracilis. Slender. 

Grain. See Caryopsis. 

Gramineous (-eus). Relating to grass or 
grain-bearing plants. 

Granular (-aris), Granulose  (-osus). 
Composed of small grains or Granulcs. 

Granulate (-atus), Granuliferus. Bear- 
ing grains or grain-like bodies. 

Gravéolens. Unpleasantlystrong-scented. 

Griseus. Gray or bluish-gray. 

Grumous (Grumosus). Consisting of 
clustered grains. 

Guttate (-atus). Spotted as if by drops. 

Gymnanthous (-us). Naked flowered. 

Gymnes. Greek for naked; used in com- 
pounds such as 

Gymnocarpous (-us). 


Pertaining to or re- 


Naked-fruited. 


414 


Gymnospermia. A Linnean artificial 
order of Didynamia, in which the 
nutlets resulting from four divisions 
of an ovary were taken for naked 
seeds; 337. 

Gymnesperms, Gymnosperme. A sub- 
class of naked-seeded plants; 268, 344. 

Gymnospermous (-us). Naked-seeded, as 
opposed to Angiospermous. 

Gynandria. A Linnean class, character- 
ized by the flower being 

Gynandrous. Stamens borne on (adnate 
to) the pistil, even to the style or 
stigma; 251, 335. 

Gynobase (Gynobasis). An enlargemeut 
or production of the torus on which 
the gynecium rests or is somewhat 
elevated; 212. 

Gyno-diecious. _ Dicecious with some 
flowers hermaphrodite and others pis- 
tillate only; 191. 

Gynacium. The pistil or collective pis- 
tils of a flower; the female portion of 
a flower as a whole; 165. 

Gynophore ( Gynéphorum). 
a pistil; 212. 

Gynostegium. A sheath or covering of 
the gvneecium, of whatever nature. 

Gynostemium. The column of an Orchid, 
consisting of andreecium and summit 
of the gvneecium combined. 

Gyrate (-atus). Curved into a circle, or 
taking a circular course. 

Gyrose (-osus). Curved backward and 
forward in turus. 


The stipe of 


Habit (Habitus). 
ance of a plant. 

Habitat. Habitation; the geographical 
limits or station; 366. 

Hematitic (-icus). Brown-red. 

Hairs. Outgrowths of the epidermis, 
consisting of single elongated cells, or 
of a row of cells. 

Hairy. Descriptively applied to pilosity 
or pubescence, in which the hairs are 
separately distinguishable. 

Hulbert- or Halberd-shaped. 
tate. 

Halved. See Diinidiate; with one half 
absent or appearing to be so. 

Humate (-atus). Hooked at the tip. 

Hamulate or Hamulose (-osus). Dimin- 
utive of Hamate. 

Haplos. In Greek compounds, simple 
or simply, as 

Haplopetalus (-us). 
of petals. 


The general appear- 


See Has- 


With only one row 


} 


GLOSSARY. 


Haplostemonous (-us). With .a single 
series of stamens; 177. 

Hastate (-alus),  Hastilis. Halberd- 
shaped, like the head of a halberd, 
v7. €. sagittate, but the basal lobes di- 
rected outward or at right angles to 
the midrib of the leaf; 96. 

Head. The form of inflorescence termed 
Capitulum, viz. a cluster of sessile 
flowers ona very short axis and centvi- 
petal in evolution; 147. 

Heart-shaped. Ovate with a sinus at 
base; 96. 

Heart-wood. The older and matured 
wood of an exogenous stem; 80. 

Hcbetate (-atus). Having a dull or blunt 
and soft point. 

felicoid (-oideus), Helicoidal. Coiled into 
a helix, or like a snail-shell. In true 
helicoid inflorescence, the flowers are 
all in a single row; 155, 157. 

Helmet. See Galea. 

Helwolus. Dull and grayish yellow. 

Hemi. Walf or halved; in Greek com- 
pounds, such as 

Hemi-anatropous. Half anatropous. 

Hemicarp (-arpium). Half or one carpel 
of a Cremocarp. 

Hemitropous (-us). Same as amphitro- 
pous or half anatropous. 

Hepta, The Greck numeral seven. 

Heptagynia. A Linnean artificial order, 
having seven styles or distinct car- 
pels; 337. ; 

Heptamerous (-us). Of seven members. 

Heptandria. The Linnzan 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. 

Herbarium, Herbal. A collection of 
dried specimens of plants, systemati- 
cally arranged; 380. 

Hercogamous (-us). Said of hermaphro- 
dite flowers when some structural ob- 
stacle prevents autogamy. 

Hermaphrodite (-itus). Of both sexes; 
191. 

Hesperidium. A hard-rinded berry, like 
an orange and lemon; 299. 


Heteracmy. Synonym of Dichogamy ; 
219 
Heteros. In Greek compounds, denotes 


(liverse or various, as 

Heterocarpous (-us, Heterocarpicus.. 
Producing more than one kind of 
fruit. 


GLOSSARY. 


Heterocephalous (-us). Bearing two 
kinds of head or capitulum. 

Tleteroclite (-itus). Anomalous in forma- 
tion, 

Ilctcrocline (-inus). Nearly same as 
Heterocephalous ; on separate recepta- 
cles. 

Heterodromous (-us). Spirals of changing 
direction. 

Heterogamous (-us). Bearing two kinds 
of flowers. 

Heterogeneous. Not uniform in kind. 

Heteroyone or Heteroyoneus. When the 
flowers are dimorphous or trimor- 
phous as respects relative length, &c., 
of stamens and pistil; 225, 234. 

Heterostyled. Same as Heterogone ; 234. 

Heteromerous. Of members not corre- 
sponding in number. 

Heteroplhyllous (-us). Having leaves of 
more than one form. 

Heterotropous (-us). Turned in more 
than one direction, or in an unusual di- 
rection; same as Amphitropous; 279. 
(Also used by Agardh for collateral 
ovules turned back to back; 282.) 

Heza. Greek numeral six; from which 
is formed 

Hexagynia. Linnean artificial order, of 
flowers with six styles or distinct car- 
pels; 337. 

Hexayynous. 
Hexagynia. 

Hexamerous (-us). Of six members; 176. 

Hexandria. Linnean class with perfect 
flowers of six stamens; 334. 

Hexandrous. Having six stamens; 249. 

Hezxapetalous (-us). Having six petals. 

Hexaphyllous. Six-leaved. 

Hexapterous (-us) Six-winged. 

Hexasepalous. Waving six sepals. 

Hexastémonous. Having six stamens. 

Hibernaculum. A winter-bud; 40. 

Hidden. Concealed from sight; as 

Hidden-veined, where the veins are in- 
visible, as in the leaves of Pinks and 
Houseleeks. 

Hiemal (-alis). Relating to winter 

Hilar ( Hilaris). Belonging to the hilum. 

Hilum. The scar or place of attachment 
of the seed; 277, 305. 

Hippocrepiform (-ormis). 
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 
bristly hairs or with briztles. 


Having the character of 


Horseshoe- 


415 


Hispidulous (-us). Minutely hispid. 

Hoary. Grayish-white with a fine and 
close pubescence. See Canescent. 

Holosericeous (-eus). Covered with fine 
and silky pubescence. 

Homocarpous(-us). With fruit all of one 
kind. 

Homodromous (-us), Homodromy. 
spirals all of uniform direction. 

Homogamous (-us). Bearing one kind 
of flowers. 

Homogeneous. All of one nature or kind. 

Homégonous or Homogone. Homomor- 
phous as respects the stamens and pis- 
til; opposed to dimorphous; 225. 

Homologue. A homologous organ or 
part. 

Homologous. Of one name or type, such 
as leaves and parts answering morpho- 
logically to leaves; 6. 

Homomdilus. Said of leaves and the like 
which are all turned in one direction. 

Homomorphous (-us). All of one form. 

Homostyled. Same as Homogone. 

Homotropous (-us). Curved or turned in 
one direction; applied also to the em- 
bryo of an anatropous seed, with rad- 
icle next the hilum; 312. 

Wood. See Cucullus. 

Hooded, Bearing or in form of a hood. 

Hornus, Hornotinus. Of the present 
year. 

Horny. Of the consistence of horn. See 
Corneus. 

Hortensis, Hortulanus. 
the garden. 

Hortus siccus. Old name of an herbarium. 

Humi. On the ground. 

Humifisus, Humistratus. 
the surface of the ground. 

Humilis. Low of stature. 

Hyaline (-inus). Transparent or trans- 
lucent. 

Hybrid. A mongrel, or cross-breed of 
two species; 321. 

Hydrophytes (Hydrophyta). 
plants. 

Hyemalis. See Hiemalis. 

Hypanthium. An enlargement or other 
development of the torus under the 
calyx; 214. 

Hypo. In Greek compounds, denotes 
under, beneath, lower. 

Hypanthodium. Same as Syconium; 
149, 303. 

Hypochilium. The basal portion of the 
labellum of an Orchid. 

Hypocraterimorphous (-us), or Hypoera- 
teriform, but the latter is a hybrid of 


With 


Pertaining to 


Spread over 


Water- 


+16 


Greek and Latin. Salverform or sal- 
ver-shaped; that is, in the form of a 
salver raised on a central support or 
stem beneath. Said of a corolla and 
the like with slender tube abruptly ex- 
panded into a flat limb; 248. 

Hiypogeous (-eus). Growing or remain- 
ing underground; 19. 

Hypogynous (-us). Under or free from 
the gyneecium or pistil; 182. 

Hypophyllous (-us). Growing on the un- 
der side of a leaf. 

Hypophyllium. An abortive leat or scale 
under another leaf, or seeming leaf, as 
in Asparagus and Ruscus. 

Hypsophylla. Answers to the German 
‘*Hochblatter,” or high leaves, those 
of the inflorescence, i. e. bracts and 
the like; 6. 

Hysteranthous (-us). With leaves pro- 
duced later than the blossoms. 


Icosandria. The Linnean class with 
twenty stamens (as the name denotes) 
or a larger number, inserted on the 
calyx; 334.  /cosandrous is the corre- 
sponding adjective; 249. 

Imberbis. Not bearded. : 

Imbricate (-atus), Imbricative. Over- 
lapping so as to ‘break joints,”’ like 
tiles or shingles on a roof; either with 
parts all in one horizontal row or cir- 
cle, as in the wstivation of a calyx or 
corolla, when at least one piece must 
be wholly external and one internal; 
or with the tips of lower parts covering 
the bases of higher ones in a succession 
of rows or spiral ranks; 135. 

Immarginate (-atus). Not margined or 
bordered. 

Immersed (-us). 
water. 

JImpari-pinnate. Pinnate with an odd 
terminal leaflet; 101. 

Inequilateral (-alis). Unequal-sided. 

Inanis. Empty, as an anther containing 
no pollen. 

Inappendiculate (-atus). 


Growing wholly under 


Not appen- 


daged. 

Incanescent. Same as Canescent. 
Incanus Hoary-white. 

Incarnate (-atus). Flesh-colored. See 
Carneus. 

Incised (-us). Cut irregularly and 


sharply; 98. 

Included (Inclusus). When the part in 
question does not protrude beyond the 
surrounding organ. 


GLOSSARY. 


Incomplete (-us). Wanting some essen- 
tial component part; 190. 

Incrassate (-atus). Thickened. 

Incubous (-us). The tip of one leaf or 
other part lying flat over the base of 
the next above it. 

Incumbent (-ens). 
upon. 

Incumbent Anther. One lying against 
the inner face of filament; 253. 

Incumbent Cotyledons, when the back 
of o:e lies against the radicle; 313. 

Jncurved (-us). Bending from without 
inward. 

Indefinite (-itus). Relates usually to 
number, this either uncertain or too 
many for easy counting. 

Indefinite Growth, 49. 

Indefinite Inflorescence, same as Inde- 
terminate; 144. ; 

/ndehiscent (-ens). Not opening by valves, 
chinks, or along regular lines; 288. 

Indeterminate. Not terminated abso- 
lutely, as the inflorescence in which 
no blossom ends the axis of the flower- 
cluster; 144, 146. 

Indigenous (-us). 
to the country. 

Individuals, 315. 


Leaning or resting 


Native and original 


Indivisus. Undivided, i. e. not cleft, 
lobed, or parted. 
Indumentum. Any hairy covering or 


pubescence which forms a coating. 

Induplicate (-atus). With edges folded 
in or turned inward. 

Indusium. The proper (often shield- 
shaped) covering of the sorus or fruit- 
cluster of a Fern. 

Induviate (-atus). Clothed with with- 
ered parts or /nduvie (clothing). 

Inequilateral. Unequal-sided; 106. 

Jnermis. Unarmed, without prickles, 
thorns, &e. 

Inferior (Inferus). Said of one organ 
when below another. In the blossom 
also in the sense of anterior; 160. An 
inferior calyx is one below the ovary, 
or free; 183. An inferior ovary is one 
with adnate or superior calyx ; 183. 

Inflated (-atus). Bladdery. 

Inflexed (-us). Bent or turned abruptly 
inward. 

Inflorescence. Mode of disposition of 
flowers ; less properly used fora flower- 
cluster itself; 141. 

Infra-azillary (-aris). Below the axil. 

Infundibuliform (-ormis), Infundibular 
(-aris). Funnelform, funnel-shaped; 
249. 


GLOSSARY. 


Innate (-atus). Borne on the apex of 
the supporting part; in an anther the 
counterpart of adnate; 252. 

Innovation (-i0). A new-formed shoot. 

Inosculating. Same as Anastomosing. 

Inseparvate, Inseparation. Terms pro- 
posed by Masters to express coales- 
cence; 181. 

Inserted (-us). 
out of. 

Insertion (-io0). 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. 

Jategérrimus. Entire in the sense of 
quite entire, z.e. the margin without 
dentation; 97. 

Inter. Between; as in Jntercellular, 

_ between the cells, &e. 
Interfoliaceous (=eus). 


Attached to or growing 


Between the 


leaves of a pair, as the stipules of | 


many Rubiacee. 
Internode (-odium). The portion of 
stem between two nodes; 6. 
Interpetiolar (-avis). Between the pet- 
ioles. 
Interruptedly pinnate. 
out a terminal leaflet. 
Intine. The inner coat of a grain of 
pollen. 
Intrafoliaceous (-eus). 
fure a leaf. 
Introflezed (-us). Same as inflexed. 
JIntrorse (-orsus). Turned inward or 
toward the axis; 253. 


Pinnate with- 


Within or be- 


Introvenius. Same as hidden-veined. 
Intruse (-usus). Pushed or projecting 
inward. 


Involicellate (-atus). Provided with a 
secondary involucre or 

Fnvolucel (-ellum). An inner or secon- 
dary involucre that of an umbellet, 
&e.; 142. 

Involicrate (-atus). 
involucre. 

Involucre (Involic: um). <A circle of 
bracts subtending a flower-cluster; 
142. 

Involute (-utus). Rolled inward: 133. 

Irregular (-aris). 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. 

Iséchrous. All of one color or hue. 

Isémerous (-us). The members of suc- 
cessive circles equal in number; 175. 


Provided with an 


27 


417 


Isostémonous (-us). The stamens just as 
many as the petals, &e. ; 177. 
Tsustemony, 196. 


Jointed. See Articulated. 

Juba. A loose panicle, with axis de- 
liquescent. 

Jugum, pl. juga. <A pair of leaflets. So 
pinnate leaves are unijugate, with a 
single pair of leaflets; bijugate, with 
two pairs; tryugate, with three pairs 
or juga, &c. Also the ridges on the 
fruit of Umbeiliferze 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 corvlla, 
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, z. 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). 
247. 

Labiatiflorous (-us). Said of certain 
Composite with bilabiate corollas. 

Labiose (-osus). Said of a polypetalous 
corolla which has the appearance of 
bilabiation. 

Labium. See Lip. 

Lacerate (Lacerus). 
if torn or lacerated. 

Lacinia. A slash; used for a slender 
lobe. 

Laciniate (-atus). 
narrow incisions. 

Lacinula. A diminutive lacinia or nar- 
row lobe. 

Lactescent (-ens). Yielding milky juice. 

Lacteus. Milk-white. 

Lacunose (-osus). Abounding in pits, 
holes, or depressions (/acune). 

Lacistrine (Lacustris). Belonging to or 
living in lakes or ponds. 


Lipped, mostly Bilabiate; 


Irregular cleft as 


Slashed; cut into 


418 


Levigate (-atus). Smooth as if polished. 

Levis (this form, and not levis, has al- 
ways been used in botany). Smooth 
in the sense of not rough. 

Lageniform (-ormis). Shaped like a 
florence flask or a gourd (the fruit of 
Lagenaria). 

Layopus. Hare-footed. 
ered with long hairs. 

Lamella. A thin plate. 

Lamellar (-aris), Lamellate (-atus), La- 
mellose (-vsus). Composed of thin 
plates or lamellz. 

Lamina. The blade or expanded part 
of a leaf, &e ; 85, 245. 

Lanate (-atus), Lanose (-osus). Bearing 
long and implexed hairs or wool 
(lana). 

Lancevlate (-atus). Shaped like a lance 
or spear-head; narrower than oblong, 
and tapering to each end, or at least 
to the apex; 95. 

Lanuginous (-vsus). Cottony or woolly ; 
clothed with soft aud implexed hairs 
or down (lanigo). 


Densely cov- 


Lappaceus. Like a bur (lappa). 
Lasianthus. Woolly-flowered. 
Latent. Undeveloped or dormant, as 


certain buds; 40. 
Lateral (-alis). Belonging to or borne 
on the sides. 


Lateritious (-ius). Of a brick-red color. 


Latex. Proper juice, milky juice, and 
the like. 

Laticiferous. Containing or conveying 
latex. 

Latiseptus. With broad partition. 

Lavender-color. Pale blue with some 
gray. 

Laxus. Loose. 


Leaf. The principal sort of appendage 
or lateral organ borne by the stem or 
axis; 85. 

Leaf-blade. The lamina of a leaf. 

Leaf-bud. A bud which develops into 
a leafy branch or its continuation; 40. 

Leaflet. A blade or separate division 
of a compound leaf; 100. 

Leaf-scar. The cicatrix left by the ar- 
ticulation and fall of a leaf; 47. 

Leafstalk. -A petiole or footstalk to a 
leaf-blade; 85, 104. 

Leathery. See Coriaceous. 

Lecus. A synonym for Corm. 

Légume (Legimen). The seed-vessel of 
Leguminose, a carpel which normally 
dehisces by both the ventral and the 
dorsal suture; 292. 

Leguminous. Pertaining to a legume, 


GLOSSARY. 


or to the order to which the legume 
gives its name. 

Lenticels (Lenticélle). 
on young bark. 

Lenticular (-aris). Lens-shaped, that is 
the shape of a lentil or a double-convex 
lens. 

Lentiginosus. Covered with minute dots 
or freckles. 

Lepal, Lepalum. A made-up word to. 
signify a stamen transformed into a 
scale, nectary, &e. 

Lepicéna. Unused name for a glume of 
Grasses. 

Lepidote (-otus). Beset with small scurfy 
scales. 

Lepis. Greek term for a scale. 

Leptos. Greek for slender; as in Lep- 
tophyllus, slender-leaved. 

Leucos. Greek for white; whence 

Leucanthus. | White-flowered. 

Leucophyllus. White-leaved, &e. 

Liber. The inner and often fibrous 
bark; 77, 81, 

Lid. See Opereulum. 

Ligneous (-eus), Lignosus. Woody. 

Ligule (Ligula). A strap or strap- 
shaped body, such the principal part 
of a ray corolla in Composite. The 
thin and scarious projection from the 
summit of the sheath of the leaf of 
Grasses, &c.; 106. Or a similar out- 
growth of the inner face of certain 
petals; 211. 

Ligulate (-atus), Liguliform (-ormis). 
Furnished with a ligule; 148, 247. 
Liguliflorous (-us). Said of the head of 
those Composite which contain only 

ligulate corol/as. 

Liliaceous. Lily-like; 246. 

Limb (Limbus). A border, 7. e. the ex- 
panded part of a gamophyllous peri- 
anth, &c., as distinguished from the 
tube and threat; 245. Sometimes the 
term is applied to the lamina or blade 
of a petal or a leaf. 

Limbate (-atus). Bordered. 

Line (Linea). 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. 

Linear (-aris). Narrow, several times 
narrower than wide, and the margins 
parallel; 95. 

Lineate (-atus). 

Lineolate (-atus). 
obscure lines. 

Lingueformis or Linguiformis, also Lin- 
culate (-atus). ‘Tongue-shaped. 


Lenticular spots 


Marked with lines. 
Marked with fine or 


GLOSSARY. 


Lip. One of the two divisions of a bila- 
biate corolla or calyx, 1. e. of a gamo- 
phyllous organ which is cleft into an 
upper (superior or posterior) and a 
lower (inferior or anterior) portion or 
lip (labium). 

Litoral or Littoral (-alis). Belonging 
to or growing on the seashore or river- 
shore. 

Livid (-idus). Pale lead-colored. 

Lobe (Lobus). Any division of an organ; 
or specially a rounded division or pro- 
jection; 98. 

Lobate (-atus) or Lobed. Divided into 
or bearing lobes; 98. 

Lobulate (-atus). Divided into small 
lobes, Lobdelets, or Lobules; 98. 

Lécellate (-atus). Divided into locelli; 
263. 

Locellus. A secondary cell; as where 
a proper cell (/oculus) of an anther or 
an ovary is divided by a partition into 
two cavities; 251, 263. 

Léculament (-entum). Sume as Loculus; 
289. 

Locular (-aris). Celied; as Dilocular, 
two-celled; ¢trilocular, three-celled, 
quadrilocular, four-celled, &c. 

Loculicidal (-idus), Loculicide. Dehis- 
cent into the cell or cavity of a peri- 
carp by the back, 7. e. through a dor- 
sal suture; 289. 

Léculus. The cell or cavity in an ovary 
or an anther. 
Loculose (-osus). 

cells. 

Lécusta. Name of a spikelet in Grasses. 

Loédicule, Lodicula. One of the small 

- seales next to the stamens in the flower 
of Grasses. 

Lomentaceous (-eus). 
sembling a 

Loment (Lomeéntum). A legume which 
is constricted or which separates into 
one-seeded articulations; 293. 

Lorate (-atus). Strap-shaped or thong- 
shaped; same as much-elongated 
linear. : 

Lucid (-idus). With a shining surface. 

Lunate (-atus). Half moon-shaped; 
crescent-shaped. 

Linulate (-atus). Diminutive of Lunate. 

Lipuline (-inus). Resembling a head of 


Partitioned off into 


Bearing or re- 


Hops. 

Lurid (-idus): Dingy-brown. 

Lusus. A ‘sport’? or variation from 
seed or bud: 319. 

Lutéolus. Yellowish; diminutive of 
Luteus. 


41%. 


Lutescent (-ens). 
faintly yellow. 

Liteus. Latin for yellow. 

Lycotropous (-us). Said of an ortho- 
tropous ovule when bent into an open 
curve or horseshoe form. 

Lyrate (-atus). Lyre-shaped; a pinnat- 
ifid form with terminal lobe large and 
rounded and one or more of the lower 
pairs small; hence Lyrately pinnate ; 
101, &e. 


Becoming yellow, or 


Macros. Greek for large or more prop- 
erly long; hence Macranthus, long- 
flowered; | Macrocephalus,  large- 
headed; Macropodus, long-footed, or 
with long stalk, &e. 

Macrospore. The larger kind of spore 
in Lycopodiacex, &c. 

Maculate (-atus). Spotted or blotched, 
i.e. with macule. 

Malpighiaceous Hairs. Those fixed by 
the middle and tapering both ways. 
Mamillate (-atus), Mamillar (-aris). 

Bearing teat-shaped jfrocesses. 

Mammeform (-ormis). Breast-shaped 

or teat-shaped; conical with rounded 


apex. 
Mammosus. Breast-shaped. 
Mancus. Deficient or wanting. 


Manicate (-atus). Said of pubescence 
so dense and interwoven that it may 
be stripped off like a sleeve. 

Marcescent (-ens), Marcidus. Withering 
without falling off; 243. 

Maryginate (-atus). Furnished with a 
margin of a distinct character or ap- 
pearance. 

Marginicidal. Dehiscent by the dis- 
junction of the united margins of car- 
pels; 290. 

Marmoratus. Marbled;- traversed by 
veins or shades of color. 

Maritime (Maritimus). Pertaining to 
the sea or seacoast. 

Mas, Masculus, Masculinus. Belonging 
to the stamens, or staminate plant, or 
flower; 191, &e. 

Masked. Sec Personate. 

Mealy. See Farinaceous. 

Medial, Median (Medianus). Belong- 
ing to the middle; in the plane of 
bract and axis; 160. 

Medificus. Fixed by the middle; 253. 

Medilla. Pith; 75. 

Medullary. Relating tothe pith J/ed- 
ullary Rays, 74; Medullary sheath, 75, 

Meiostémonous. With fewer stamens 
than petals. 


420 


Melleus. Having the taste or smell of 
honey. 
Melligo. Honey-dew. 


Membranous, Membranaceous (-eus). 
Thin and rather soft or pliable, like 
a membrane. 

Meniscoid (-oideus). 
like a meniscus. 

Mericarp (-arpium). One of the akene- 


Concayo-conyex, 


like carpels or a closed half-fruit of | 


Umbellifera ; 297. 
Mevismatic. Dividing into parts or sim- 
ilar portions. 
Merithallus. Synonym of Internode. 
Merous (-us). In Greek compounds, de- 
notes parts or members; lence Dime- 
rous, of two parts, &e. 
Mesocarp (-arpium). The middle layer 
of a pericarp; 285. 
Mesophleaum. ‘The middle or green 
bark ; 76. 
Metamorphosis, Metamorplhy, 167. 
Micropyle (-yla). The spot or point in 
the seed at which was the orifice of the 
ovule; 277, 305. 
Microspore. The smaller kind of spore 
in Lycopodiacee, &c. 
Midrib. The central or main rib; 92. 
Miniate (-atus). Vermilion-color. 
Mistus or Miztus. A cross-breed ; 321. 
Mitreform or Mitriform (-ormis). Mi- 
tre-shaped or cap-shaped. 
Monadelphia. The Linnzean class con- 
taining flowers with Monadelphous 
stamens, 7. €. those united by their 
filaments into a tube or column; 250, 
335. 
Monandria. The Linnzan class (334) 
“containing flowers with Jfonandrous, 
that is, a single stamen; 249. 
Monadnthous (-us). One-flowered. 
Moniliform (-ormis). Necklace-shaped ; 
cylindrical and with contractions at 
intervals. 
Monocarpellary. Of one carpel : 261. 
Monocarpic (-icus). Monocarpous, Mon- 
ocarpian. Only once fruiting; 33. 
Monvcephalous (-us). Bearing a single 
capitulum. 
Monochasium. A cyme with one main 
axis; 152, 155. 
Monochlamydeous (-eus). Having but 
one kind of perianth; 190, 340. 
Monvoclinous (-us). Synonym of Herma- 
phrodite. 
Monocotyl don, Monocotylédones, adj. 
Monocotyledonous (-eus). Plants or em- 
brvo with a single cotyledon; 23, 27, 
314, 339. 


GLOSSARY. 


Monolocular (-aris). One-celled. 

Monecia. Name of Linnean class (335) 
with flowers. 

Monecious (-ius), or Monvicous (-us). 
With stamens and pistils in separate 
blossoms on the same plant; 191. 

Monogamia, Name of a Linnzean arti- 
ficial order, in class Syngenesia; 337. 

Monograph. A systematic account of 
parucular genus, order, or other group; 
369. 

Monogynia. Name of a Linnean arti- 
ficial order, with solitary pistil, or 
style; 337; hence, adjectively, Mond- 
gynous ; 267. 


Mondémerous. Formed of a single mem- 
ber; 176. 
Monopetdlous (-us). Literally one- 


petalled; but always used in the 
sense of Gamopetalous, which term is 
to be preferred ; 244. 

Monophyllous (-us). One-leaved. 

Ménopode (Monopdédium), Monopodial. 
A stem of a single and continuous 
axis; 55. 

Mondplerous (-us). One-winged. 

Monepyrcnus. Containing a single stone 
or nutlet. j 

Monosepalous (-us). Equivalent to Gam- 
osepalous; but literally of a single 
sepal; 244. 

Monospcrmous (-us). 

Mi mnostichous (-us). 
rank. 

Monostylous (-us) With a single style. 

Monosymmetrical. That which ean be 
bisected into equal halves in only one 
plane; 175. 

Monétocous (-us). Bearing progeny 
(fruiting) only once, as annuals and 
biennials ; 33. 

Monster (Monstrum). A monstrosity, 
or unnatural development. 

Morphology, 5. 

Moschate (-atus). Exhaling the odor of 
musk. 

Mueilaginous, Mueilaginosus. Slimy; 
of the consistence or appearance of 
mucilage. 

Mucro, Mucronation. A short and 
abrupt small tip to a leaf, &e. 

Micronate {-atus). Tipped with a mu- 
cro; 97. 

Mucrénulate (-atus). 
nate. 

Mule. A hybrid or cross-breed. 

Multicipital ( Multiceps). Many-headed ; 
many shoots or stems from the erown 
of one root. 


One-seeded. 
In a single vertical 


Minutely muero- 


GLOSSARY. 


Multifarious (-us). Many-ranked, as 
leaves in several vertical ranks 

Miultifid (-idus). Cleft into many lobes 
or segments. 

Multiflorous (-us). 

Multijigate (Multijugus). 
pairs or juga. 

Multilocular (-aris). 
several-celled. 

Multiparous. Many-bearing; said of a 
several-branched cyme; 152, 158. 

Multiple fruits. The tructification of a 
flower-cluster when coufluent into one 
mass; 301. 

Multiplication. Same as Augmentation ; 
179, 200. 

Multiserial (-alis), Multisertate (-atus). 
In several series. 

Miurvicate (-atus). Rough with short and 
firm excresceices. 

Muriculate (-atus). Minutely muricate. 

Museariformis. Vly-brush-shaped. 

Musciform (-ormis). Moss-like in ap- 
pearaiice. 

Muscoloyy. The botany of Mosses. But 
is a hybrid word, and is replaced by 


Many-flowered. 
In many 


Many-celled or 


Bryology. 

Miticous (-us). Pointless, blunt, awn- 
less. 

Myeclium. The filamentous vegetative 


growth of a Fungus. 
Mycology, Mycetology. The botany of 
Fungi. q 
Mycropyle. Micropyle misspelled. 


Naked. Wanting some usual covering ; 
as flowers without perianth, ovules 
without coats, seeds not in a pericarp, 
buds without seales. 

Napiform (-ormis). Turnip-shaped ; 31. 

Nanus. Dwarf. 

Natant (-ans). 
under water. 

Navieular (-aris). 
as Cymbiform. 

Nebulose (-osus). Clouded or misty. 

Neck. See Collum. 

Necklace-shaped. See Moniliform. 

Nectar. The sweetish secretion by va- 
rious parts of the blossom from which 
bees make honey. 

Nectary (Necturium). The place or 
thing in which nectar is secreted: 
formerly applied also to any anoma- 
lous part or appendage of a flower, 
whether known to secrete honey or 
not; especially to the hollow spurs of 
a Violet, Larkspur. Columbine, and 
the like. 


Floating or swimming 


Boat-shaped. Same 


421 


Nectariferous (-us). Nectar-bearing. 
Needle-shaped. See Acerose. 


Nemorosus, Nemoralis. Inhabiting 
groves. 
Nervation. Same as Venation, or un- 


branched venation. 

Nerve (Nereus). In botany, this is a 
simple or unbranched vein, or a slen- 
der rib. 

Nerved, Nervose (-osus), Nervate (-atus). 


Having ueryes in the botanical 
sense. 

Nervulose (-osus). Diminutive of ner- 
vose. 

Netled. Same as Reticulated; Netted- 


veined; 92. 

Neurose (-osus). Same as Nervose. 
Neura being the Greek for nerve. 
Neuter, Neutral. Sexless; as a flower 
which has neither stamen nor pistil; 

191, 195. 

Niger. Black or blackish. 

Nigricuns. Turning black or verging 
to black. 

Nitidus. Smooth and shining. 

Nicalis. Growing in or near snow. 

Niveus. Suow-white. 

Nodding. Uanging down. 

Node (Nodus). Literally a knot; the 
portion of a stem which normally 
bears a leaf or whorl of leaves; 6. 

Nedose (-osus). Kuotty or knobby. 

Nodulose (-osus). Diminutive of Nodose. 

Nomenclature, 3, 345. 

Normal (-alis). According to rule; 
agreeing with type. 

Notate (-atus). Marked by spots or lines. 

Nothus. Yalse or bastard. 

Notorhizal (-izus). Synonym of incum- 
bent, as applied to the embryo of 
Cruciferz. 

Nuciform (-ormis). Nut-like in shape. 

Nucleus. A kernel of an ovule, seed, 
&e.; 277. A soft solid interior part 
of a vegetable cell in the early condi- 
tion; 309. 

Nuculanium. Name given by Richard 
to a drupaceous or baccate fruit con- 
taining more than one stone or stony 
seed; adopted by Lindley for a supe- 
rior stony-seeded berry, such as a 
grape. 

Nucule (-ula). A diminutive nut or 
stone; same as Nutlet; 296. 

Nucum+ntaceus (-eus). Nut-like in char- 
acter. 

Nudicaulis. Naked-stamened ; stem not 
leafy. 


Nudus. Naked, in its various senses. 


422 


Numerous (Numerosus). Used in the 
sense of indefinite in number. 

Nut (Nuzx). A hard and indehiscent 
one-seeded pericarp resulting from a 
compound ovary; 295. 

Nutlet. A diminutive Nut. See Nucule. 

Nutant (-ans). See Nodding. 


Ob. Over against; as a prefix denotes 
inversely or oppositely, as 

Obcompressed (-us). Flattened the other 
way, antero-posteriorly instead of lat- 
erally. 

Obconical (-icus). Conical, but attached 
at the apex. 

Vbcordate (-atus). Inverted _heart- 
shaped, the notch at the apex; 97. 
Obdiplostemonous (-us), Obdiplostemony. 
When the stamens are double the 
number of the petals, but the outer 

series opposite the latter; 198 

Obimbricate. Imbricated or successively 
overlapping downward. 

Oblanceolute. Lanceolate but tapering 
toward the base more than toward the 
apex ; 95. 

Oblique ( Obliquus). 
slanting. 

Oblong ( Oblongus). Considerably longer 
than broad and with nearly parallel 
sides; 95. 

Obovate (-atus). Ovate with the broader 
end toward the apex; 95. 

Oboroid Solid obovate. 

Obtectus. Covered by something. 

Obtuse (-usus). Blunt or rounded at the 
extremity ; 97. 

Obtusiusculus. Somewhat obtuse ; dimin- 
utive of obtuse. 

Obvallatus. Guarded on all sides or sur- 
rounded as if walled in. 

Obverse, Obversely. Same as the prefixOb. 

Obvolute (-utus). A modification of Con- 
volute; 138, 139 

Ocellate (-atus). With a circular patch 
or a ring of color. 

Ochraceous (-eus). Ochre color; light 

_ yellow with a tinge of brown. 

Ochrea, Ocrea. A leggin-shaped or 
tubular stipule or rather combined 
pair of stipules; 106. 

Ochreate, Ocreate (-atus). Furnished 
with ochrea or sheaths; 106. 

Ochroleucous (-us). Yellowish-white or 
between white and yellow. 

Octo. Eight. In composition gives such 
terms as the following. 

Octacynia. Linnean artificial order with 
Octagynous (eight-sty led) flowers ; 337. 


Unequal-sided or 


GLOSSARY. 


Octamerous. 
the circle. 

Octandria. The Linnean class with 
Octandrous, i. e. eight-stamened flow- 
ers; 249, 334. 

Octéni. In eights. 

Octopetalous (-us). Eight-petalled. ~ 

Octosepalous. With eight sepals. 

Octostichous (-us). In eight vertical 
ranks; 124. 

Oculate (-atus). Same as Occllate. 

Officinal (-alis). Used in medicine or 
the arts, therefore in the shops. 

Offset. A short lateral shoot for propa- 
gation; 53. 

Oides, Oideus, -odes and -ides. Greek 
for likeness, used as terminations, in- 
dicate similarity to; as Dianthoides, 
resembling a Pink. 

Oleraccouz (-cus). Esculent in the way 
_of a pot-herb. 

Oligos. Greek for few; in compounds 
giving such terms as 

Oligandrous (-us). With few stamens. 

Oliganthous (-us). Few-flowered. 

Oliyémerous (-us). Of few members. 

Oligospérmous (-us). Few-seeded. 

Olivaceus (-eus). Olive-green. 

Omphalodium. A mark (navel) on the 
hilum of a seed, through which passed 
vessels to the chalaza or rhaphe. 

One-sided. Fither turned to one side, 
or with parts all turned one way, or 
unequal-sided. : 

Oophovidium. The spore-case for the 
larger spores in Selaginella, &c. 

Opaque (Opdcus). Mostly used in the 
sense of not shining or dull. 

Operculate (-atus). Furnished with an 
Operculum or lid. 

Operculum. <A lid; a top which sepa- 
rates by a transverse line of separation 
as does that of a pyxis. 

Opposite (-itus). Set against; as leaves 
over against each other when there 
are two on one node; or one part be- 
fore another, as a stamen before a 
petal; 6, 120, 178. 

Oppositifolius (-ius). Placed opposite a 
leaf, as is a tendril or peduncle in 
Vitis, &e. 

Oppositipetalous (-us). 


Composed eight parts in 


Placed before a- 


petal. 

Oppositisepalous (-us). Situated before 
a sepal. 

Orbicular (-arvis), Orbiculate (-atus). 


Said of a flat body with a circular out- 
line; 95. 
Orchidacevus, 246. 


GLOSSARY. 


Order (Ordo). Group between genus (or 
tribe) and class; 328. 

Ordinal. Relating to orders. 

Organdgeny (Organogenesis). The for- 
mation or early development of or- 
gans ; 2. 

Organéography, Oryandlogy. The study 
of organs and their relations; 2. 

Organs of Vegetation, 11. 

Orgyalis. Six feet high, or of the height 
of a man. : 

Ornithéphilous. Said of flowers which 
are habitually fecundated by pollen 
brought by birds; 217. 

Orthoploceus. Said of an embryo when 
incumbent cotyledons «re folded 
around the radicle, as in Mustard 

Orthos, Greek for straight; whence com- 
pounds such as the following 


Orthostichies. Vertical ranks; 121. 
Orthostichous. Straight-rauked. 


Orthétropous (-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 stene, same as Pyrena. 

Ostidlate (-atus). Furnished with a small 
orifice or little door ( Ostiolum). 

Outgrowths, 209. 

Oval (Ovalis). 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 (Ovvideus). Used either for solid 
ovate or solid oval, more properly for 
the latter. 

Ovulate (-atus), Ovuliferous. 
ovules. 

Ovule, Ovuluim. The body in the flower 
which becomes a seed; 166, 276. 


Bearing 


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 Compositx; also 


| 


i 


423 


an inner bract or glume in Grasses; 
142. 

Paleaceous (-eus). Chafiy ; furnished 
with pale; 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. 
Paleolate (-atus). 

ole. 

Palets (Pales of some Evglish botanists). 
Same as Paleze; 142. 

Palmaris. A palms breadth or length; 
z. 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-). 
the palmate manner. 

Pulmately veined, 93 

Palmatifid (-idus), Palmatilobate, Pal- 
matisect). Palmately cleft, lobed, or 
divided. 

Palminerved. Palmately nerved; 93. 
Paludose (-osus), Palustiine (Palustris 
or Paluster). Inhabiting marshes. 
Pandurate (-atus),Panduriform (-ormis). 

See Fiddle-shaped. 

Panicle (Panicula). A loose compound 
flower-c luster, 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. 

Papillar (-aris), Papillose (-osus), Papil- 
late (-atus). Bearing or resembling 
papille, minute nipple-shaped projec- 
tions. 

Pappiferous (-us), 
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 (-us), Papulose (-osus). 
Covered w.th Papule, or small pim- 
ples 


Furnished with pale- 


In 


Puppose  (-osus). 


424 GLOSSARY. 


Paracarpium. Unused name given to 
an abortive pistil or carpel. 

Puracorolla. A crown or internal ap- 
pendage or deduplication of a corolla. 

Parallel-nerved, veined, &e. Same as 
Nerved; 91. 

Parapetalous (-us). Said of stamers, 
&c., which stand at each side of a 
petal; 178, 201. 

Pardphysis, pl. Paraphyses. Jointed 
thread-like bodies, of no known func- 
tion, accompanying the archegonia of 
Mosses. 

Parasitic (-icus). Growing on or in and 
living upon another plant or even ani- 
mal; 36. 

Purastémon. Name rarely applied to an 
abortive stamen or body in place of or 
accessory to a stamen; same as Sta- 
minodium. 

Parastichies Secondary spirals in phyl- 
lotaxy; 127. 

Parenchyma. Common or soft cellular 
tissue. 

Parenchymatous. Of the nature of or 
composed of parenchyma. 

Paries, pl. parietes. The wall of any 
organ. 

Parictal (Parietalis). Borne on or re- 
lated to the wall; 265. 

Paripinnate (-atus). Even-pinnate ; same 
as abruptly pinnate; 101. 

Parted, Partite (-itus). Cleft nearly 
but not guite to base; 98. 

Parthenogénesis, Parthenogeny. Pro- 
duction of seed without the interven- 
tion of pollen; 285. 

Partial (Partials). Secondary, as Par- 
tial involucre (142), peduncle (143), 
petiole (105), umbel (150), &c. 

Partible (Partibilis). At length sepa- 
rating or easily to be separated. 

Partition. In one sense a separated por- 
tion or segment; in another and the 
more usual, a wall or dissepiment. 

Patelliform (-ormis). Disk-shaped, cir- 
cular with a rim, of the form of the 
patella or kneepan. 

Patent (Patens) Spreading ;_ either 
widely open or diverging widely from 
an axis. 

Patentissimus. Superlative of Patens; 
extremely spreading. 

Patulous (-us) Slightly or moderately 
spreading. 

Pauciflorous (-us). Few-flowered. 

Paucifolius. Few-leaved. 

Petr-shaped. Obovoid or obeonical with 
more tapering base. 


Peéctinate (-atus). Pinnatifid with nar- 
row and closely set segments, like 
comb-teeth. 

Pedilis A toot long or high. 

Pedate (-atus). Palmately divided or 
parted with the lateral divisions two- 
cleft; resembling a bird's foot. 

Pedatipartitus, -lobatus, -sectus, &e. Pe- 
dately parted, lobed, divided, &e. 

Pedicel (-clius). An ultimate flower- 
stalk or its division; the support of a 
single flower; 143. 

Pedicellate (-atus). Pedicelled, borne 
on a pedicel. 

Pediculus. Name sometimes used for 
Pedicel. 

Pedunele (Pedinculus). A general 
flower-stalk, supporting either a clus- 
ter or a solitary flower; in the latter 
case,the cluster may be regarded as 
reduced to a single blossom; 143. 

Pedinculate (-atus). Peduncled, borne 
on a foot-stalk. 

Pelovia. An irregular flower become 
regular by a monstrous development 
of complementary irregularities ; 186. 

Peltate (-atus), LPeltiforvm  (-ormis). 
Shield-form ; target-shaped; a plane 
body attached by its lower surface 
(instead of margin or base) to a stalk; 
96, 107. 

Peltinerved (-ius) Radiately-nerved or 
ribbed all round the circle. 

Pelviform (-vrmis). Basin-shaped ; shal- 
low cup-shaped. 


Pendent (-cns). Hanging on its stalk or 


support. 

Pendulus (-us), Pendulinus. Hanging 
more or less, as if from weakness of 
the support. 

Penicillate (-atus), Penicilliform (-ormis). 
Pencil-shaped, the pencil (penicillum) 
being a brush or tuft of hairs. 

Pennute (-atus). Same as Pinnate. 

Penniform (-ormis). In the form of a 
feather or its plume. 

Penninerved (-ervius). Same as pin- 
nately nerved or veined; 93. 

Penta. Greek for five; gives compounds 
such as 

Pentacarpellary. Composed of five car- 
pels; 261. 

Pentachenium. Name of a pentacarpel- 
lary fruit otherwise like a cremocarp. 
Pentadeélphous (-ws). With stamens in 

five clusters; 250. 

Pentacynia. Linnean. artificial order 
characterized by Pentagynous, 7. € 
five-styled flowers; 337. 


GLOSSARY. 


Pentamerous (-us). Composed of five 
members in a circle; 176. 

Pentandria. The Linnean class with 
Pentandrous, i. e. tive-stamened flow- 
ers; 249, 534. 

Pentap:talous (-us). Five-petalled; 244. 

Pentaphyllous \-us). Vive-leaved: 243. 

Pentapterous (-us). Five-winged. 

Pentasepalous(-us). Of five sepals; 244. 

Pentastichous(-us). In tive vertical ranks ; 
123. 

Pepo, Peponida, Peponium. 
fruit; 298. Z 

Perennial (Perennis, Perennans). Last- 
ing year after year; 32. 

Perfect (Perfectus). Said of a flower 
which is hermaphrodite. 

Perfoliate (-atus). Where a stem seem- 
ingly passes through a leaf; 167. 

Perforate (-atus). 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 (-arpium). The  fructitied 
evary; 286. 

Pericarpic (-icus). 
carp. 

Perichetial (-ialis). Relating to the 
Perichetium, a set of bracts around 
the fruit-stalk in Mosses. 

Pericladium. The sheathing base of a 
leaf when it expands and surrounds 
the supporting branch. 

Periclinium. Involucre of the capitu- 
lum of Composite; 148. 

Péviderm (-ermu or -ermis) 
or Epiphlceum. 


A gourd- 


Relating to the peri- 


Outer bark 


Pévigone, Perigénium. 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 Cyperacez. 
Perigynous (-us} Literally around the 
ovary: said of organs which are ad- 
nate to the perianth, or to this as con- 
nate with the lower part of the pistil; 
182. 
Peripétalous (-us). Around the petals. 
Peripheric (-icus). Of or belonging to the 
circumference ; as of an embryo coiled 
round the outside of the albuinen. 
Periphoranthium. Synonym of the 
involucre of Composite; 148. 


425 


Peripterus. Surrounded by a wing or 
thin border. 

Perisperm (-ermium). The albumen of 
the seed, at least the exterior and or- 
dinary albumen; 14, 310. 

Peristome (Peristoma or Peristémium). 
The fringe or other structure surround- 
ing the orifice (stoma) of a Moss. 

Peritropous (-us) or Pevitrepal. 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. 

Peérsonate (-atus) Masked, as when a 
bilabiate corolia has a prominent pal- 
ate; 248. 

Pervious (-ius). 
way. 

Pertuse (-usus). Having slits or holes. 

Perula, pl. Perule. Scales of leaf-buds 
and the like; 40. 

Perulate (-atus). Furnished with peru- 
le or scales. 

Pes, gen. pedis. A foot. Hence in 
Latin compounds Lonyg/pes, long- 
stalked, Brevipes, short-sialked, &e. 

Petal (Petalum). <A corolla-leaf; 165. 

Petaline (-inus), Petalvid (-oideus). 
Petal-like, or relating to petals; 118. 

Petalody. Name for the metamorphosis 
of other organs (such as stamens) into 
petals; 174. 

Petiolar (-aris). 
to a petiole. 

Petiolate (-atus), Petioled. 
petiole. 

Peétivle (Petiolus). 
leaf; 85, 104. 
Petidlulate (-atus), 

Having a 

Pétiolule (Petidlulus). 
leaflet; 105. 

Petreus. Growing among rocks. 

Petrosus. Growing in stony places. 

Phenogams, Phenogamia, Phenoga- 
mous plants. Plants sexually propa- 
gating by flowers, of which the essen- 
tial organs are stamens and pistil; 3, 
334, 340, 344. 

Phalanges, sing. Phalanz. The bundles 
of stamens in diadelphous or polyadel- 
phous flowers. 

Phanerogams, Phanerogamia, &e. See 
Phenogams, «ce. 


With an open passage- 


Borne on or relating 
Having a 
The footstalk of a 
Petiolular 


(-a7ris). 


A footstalk of a 


Phleum. Greek name for bark. 

Pheniceus. Deep red with some sear- 
let. 

Phoranthium. A name for the recep- 


426 


tacle of the capitulum in Composite ; 
148. 

Phycology. The botany of Algze. 

Phylla. Leaves in Greek; combined 
with Greek numerals, forming such 
terms as Diphyllous, Triphyllous, &c., 
to Polyphyllous. : 

Phyllocladium. A branch assuming the 
function of foliage; 65. 

Phyllodineous (-eus). Relating to a 

Phyliidium. A petiole usurping the 
form and function of a leaf-blade; 
110. 

Phyllody, Phyllomorphy. Names for 
the transformation or metamorphosis 
of floral organs into leaves; 174. 

Phyllotaxis, Phyllotary. Leaf-arrange- 
ment; 119. 

Phyllomania. The unusual or abnormal 
production of leaves. 

Phyllophore (-orum). The budding sum- 
mit of a stem on which leaves are de- 
veloping. 

Phyllum. 
Phylla. 

Phyllome, Phylloma. An assemblage 
of leaves, or of incipient leaves in a 
bud. Also recentiy used by German 
botanists for leaf generically or poten- 
tially, that which answers to a leaf; 6. 

Phytography. Botany as relates to the de- 
scription and illustration of plants ; 345. 

Phy dlogy. Synonym of Botany. 

Phiytomer, pl. Phytomera. Plant-ele- 
ments in morphology ; same as 

Phyton. Greek name for plant; has 
been used in the sense of plant-ele- 
ment, or plant-unit; 7. 

Phytotomy. Same as Vegetable Anatomy 
or Histology; 2. 

Piceus. Pitch-black or brownish-black. 

Pictus. Painted, or rather as if painted. 

Pileate (-atus), Pileiformis. Having the 
form of a cap or Pileus. 

Pileorhiza. The root-cap. 

Piliferous (-us). Bearing or tipped with 
hairs (pili). 

Pilusciusculus. Slightly hairy. 

Pilose (-osus). Hairy, in general with 
any sort of pilosity ; in particular with 
soft and distinct hairs. 

Pinna. One of the primary divisions of 
a pinnate leaf, either simply pinnate, 
when it is a leaflet, or a partial petiole 
or rhachis with the leaflets when the 
leaf is bipinnate; 104 

Pinnate (-atus). When leaflets are 
arranged along each side of a com- 
mon petiole ; 100. 


Greek for leaf; 6, 85. See 


GLOSSARY. 


Pinnately cleft, lobed, parted, &e.; 99. 

Pinnately veined. Yeather-veined; 93. 

Pinnatifid (-idus). Pinnately cleft. - 

Pinnatilobatus, Pinnaidobus. Pinnately 
lobed. 

Pinnatipartitus. Pinnately parted. 

Pinnatisictus. Pinnately divided quite 
down to the rhachis. 

Pinnule (Pinnula). One of the pinnately 
disposed divisions of a pinna; a sec- 
ondary pinna; 104. 

Pisiform (-ormis). Pea-shaped; resem- 
bling a pea. 

Pistil (Pistiilum). The female organ‘ of 
a flower, consisting of ovary, style. 
aud stigma, or at least of ovary and 
stigma; 302, 259. 

Pistillate (-atus), Pistilliferous. Said of 
a plant or a blossom provided with 
pistil, most properly for one having 
pistilonly; 191. 

Pistillidium. One of the names of the 
analogue of pistil in Mosses, &e. 

Pistillody. Name for the metamorpkosis 
of other organs into carpels; 174. 

Pitcher. See Ascidium. A tubular or 
cup-shaped leaf, which usually holds 
some liquid; 111. 

Pith. A ceutral cellular part of a stem, 
especially of an exogenous stem; 75. 


Pitted. Marked with small depressions 
or pits. 
Placenta. That in the ovary which 


bears the ovules, sometimes the mere 
united margins of the carpel-leaves, 
somctimes a thickening or enlarge- 
ment of them, or even of some other 
part of the ovary; 261. 

Placentation (-iv). The disposition of 
the placentz. 

Placentiform (-ormis). Quoit-shaped, 
or in form like a flat cake. 

Plaited. See Plicate. 

Plane (Planus). With flat surface or 
surfaces. 

Platys. Greek for wide, in such com- 
pounds as Platyphyllus, broad-leaved, 
&e. ; 

Pleios. Greek for full, used in com- 
pounds for several or many; as Pleto- 
phyllous, several-leaved, &e. Simi- 
larly Pleistos for a great many. 

Pleiockdsium. A several-rayed cyme; 
152, 155. 

Plenus. Full. Flos plenus is what gar- 
deners call a ‘‘ double flower,’ that is 
one in which the petals or other flower- 
leaves are abnormally multiplied. 

Pleur-nchymat. Same as woody tissue. 


GLOSSARY. 427 


Pleurorhizal (-us). Embryo with radicle 
against one edge of the cotyledons; 
z. e. the latter accumbent. 

Plicate (-atus), Plicativus. Folded into 
plaits (plice), usually lengthwise; 133, 
139. 

Plimbeus. Lead-colored; 
with some metallic lustre. 

Plumose (-osus). Feathered; when bris- 
tles, &ec., have fine hairs on each side 
like the plume of a feather, as the pap- 
pus of Thistles. 

Plumule (Plumula). 
ing point of the embryo above the 
cotyledons; 17. 

Plures. Many or several; used as a 
prefix in Latin words, such as Pluri- 
florous (-us), several-tlowered ; Pluri- 
loculur (-aris), several-celled ; Pluri- 
Joliolate, with several leaflets, &c., 
Plurijugate, in several pairs, &e. 

Péculiform (-ornis). In the shape of a 
drinking-cup or goblet. 

Pod. A dry and several-seeded dehis- 
cent fruit; strictly a Leguine or a 
Silique ; 288, 292. 

Podium, Podus. A footstalk, stipe, or 
other such support; used only in Greek 
compounds, as Podocephalus, head 
pedunculate; Podocarpus, fruit stipi- 
tate; or as a suffix, in such words as 
Leptopodus, slender-stalked; Brachy- 
podus, short-stalked, &c. 

Podetium. Any stalk-like elevation. 

Podogynium. Same as Gynophore. 

Podosperm (-ermium). The stalk of a 
seed; 276, 305. 

Pogon. Greek fora beard ; enters into 
various compound words. 

Polémbryony. See Polyémbryory. 

Politus. Polished; applied to a smooth 
and shining surface. 

Pointless. Same as Muticous. 
Pointletted. Minutely pointed; same as 
apiculate or as minutely acuminate. 
Pollen, Pollen-grains. The fecandating 
grains or cells contained in the anther; 

165, 256. 

Pollen-tube. The slender tube which 
begins as a protrusion of the inver 
coat of a pollen-grain, and elongates 
by growth, at least when in contact 
with the stigma; 258. 

Pollicaris. An inch long; the length 
of the terminal joint of the thumb, 
pollex. 

Poliniferous (-us). Pollen-bearing. 

Pollinium. A mass of pollen-grains 
more or less coherent ; 257, 230. 


dull gray 


The bud or grow- 


| 


Pollinated (-1tus)}. Said of a stigma 
when supplied with pollen. 

Poly. In Greek compounds, denotes 
numerous ; as in 

Polyadelphia. Name of a Linnzean ar- 
titicial order with stamens Polyadel- 


phous, or in several phalanges or 
brotherhoods; 250, 335. 
Polyandria. Name of a Linnean class 


with flowers Polyandrous, or having, 
an indefinite number of stamens; 249. 
334. 

Polydnthous (-us). Many-flowered;. in 
the Latin form same as multiflorous. 
Polycarpellary. Of many carpels; 261. 
Polycarpic (-icus). Fruiting many times 
or indefinitely; DeCandolle’s name 

for a perennial herb; 33. 

Polycephalous (-alus). Consisting of or 
bearing many heads, eupitula. 

Polycoccus. Of several cocci. 

Polycotyledonous (-eus oF es). 
several cotyledons; 22, 314. 

Polygamia. Name of a Linnzean class 
having Polggaimous flowers, i. e. some 
hermaphrodite, some unisexual ; 191, 
335. Also of Linnean orders of Syn- 
genesia; 337. 

Polygynia. Name of a Linnzean artifi- 
cial order with flowers Polygynous, 
7. e. containing numerous carpels; 
261, 337. 

Polymerous (-us). Of numerous mem- 
bers to each series or circle. 

Polymorphous (-us). Of several or vari- 
ous forms. 

Polypetalous (-us). 
petals; 244. : 

Polyphore (-drium). A torus which 
bears many pistils, as that of a straw- 
berry or raspberry} y- 

Polyphyllous (- -us). Many-leaved; 244. 

DP lysepalous (-us). Of separate sepals; 
244 

Polyspérmous (-us). 

Polystemonous (-us). 
mens. 

Polystachyus. Bearing many spikes. 

Polystylous (-us). Bearing many styles. 

Polysymmetrical. That which can be 
divided into similar halves in several 
or more than one plane; 175. 

Polytocous (-us). Bearing progeny (fruit- 
ing) many times, 7. c. year after year; 

Pome (Pomum). Kind of fruit of which 
the apple is the type; 298. 

Pomeridianus. In the afternoon. 

Pomiferous (-us)., Pome-bearing. __. 


Having 


Having separate 


Many-seeded. 
With many sta- 


428 


Pomology. A treatise on or the subject 
of fruits considered as esculent. 

Porouse (-osus), Porous. Pierced with 
small holes or pores. 

Posterior. In an axillary flower is the 
side next the axis of inflorescence; 
160. 

Posticous (-us). On the posterior side, 
which in a flower is that next the axis 
of inflorescence: an adnate anther is 
posticous when on the outer side of 
the filament, 7. e. when it faces the 
petals; 253. 

Pouch. See Silicle. 

Precot. Appearing or developing early 

Prefloration. Same as Hstivation; 132. 

Prefoliation. Same as Vernation; 132. 

Premorse (-orsus). With end as it were 
bitten off. 

Prasinus. Grass green. 

Pratensis. Growing in meadows. 

Prickly. Armed with Prickles (56), 
which are outgrowths of the bark or 
rind. 

Primine. Outer coat of the ovule; 277. 

Primordial (-ialis). The first in order 
of appearance. Primordial leaves are 
those of the plumule. 

Prismatic (-icus). Prism-shaped, with 
flat faces separated by angles. 

Procerus. Very tall. 


Process (Proccssus). Any projecting 
appendage. 


Procumbent (-ens). 
ground; 53. 

Productus. Produced, 7. e. extended or 
prolonged into. 

Pro-embryo, 284. 

Proles. Progeny; sometimes used for 
race; 32i. 

Proiiferous (Prélifer, Proliferus). Bear- 
ing progeny, in the way of offshoots. 

Proliferation or 

Prolification is usually taken as the 
production by one organ of something 
different, such as the development of 
buds and plantlets on leaves, of leafy 
shoots in place of flowers, &c.: 73. 

Proligerous (-us). Same as Proliferous. 

Prone (Pronus). Lying flat, especially 
face downward. 

Propaculum, Propagulum. Name of a 
shoot, such as a runner or sucker which 
may serve for propagation. 

Propdgines. Same as Bulblets. 

Prophylla. Primary leaves, as the first 
leaves of a b-anch or axis. 

Prosenchyma. Plant-tissue consisting of 
lengthened, tubular, or fasiform cells. 


Lying along the 


GLOSSARY. . 


Prostrate (-atus). 
the ground; 53. 

Protos. Greek for first; used in various 
compounds, such as 

Protundrous, Protand:y. 
androus. 

Proterdndrous, also Protandrous, Pro- 
terandry. When the anthers of a 
flower are in alithesis earlier than the 
stigma; 219, 220. 

Proteranthous (-us). 
precedes leafing. 

Proterogynous, Proterogyny, or Pro- 
togynous, Protogyny. When the 
stigma is ready for its functious ear- 
lier than the anthers of the same 
blossom; 219. 

Protophytes, Pretophyta. 
the supposed first plants. 

Protoplasm, Protoplasma. The forma- 
tive organic material of plauts and 
animals, in its living state. 

Pruinate (-atus), Pruinose (-osus). As if 
frosted over with a bloom or powder. 

Pseudos. Greek for false, a prefix in 
various compounds, as Pseudo-mono- 
cotyledonous ; 26. 

Pseudo-bulb. A thickened and bulb- 
like internode in epiphytal orchids; a 
corm. 

Pseudocarp (-arpium). The principal 
or accessory part of an anthocarpous 
fruit; 300. 

Pseudo-costate False-ribbed, as where 
a marginal or intramarginal vein or 
rib is formed by the confluence of thie 
true veins. 

Pseudospermium. Name given to any 
kind of «ne-seeded fruit which is inde- 
hiscent and resembles a seed, such as 
an akene, &e 

Psilos. Greek for naked or bare; as in 
Psilostachyus, with naked spike. 

Pteridium, Pteiodium. Names for the 
Key-fruit or Samara. 

Pteris. Used for wing in Greek com- 
pounds, also for a Fern. 

Pteridographia. The botany of Ferns. 

Pterocarpous (-us). Wing-fruited. 

Pteropodus. Wing-footed, t. e. petiole 
wing-inargined, &c. 

Ptyzis. Greek name for folding, as of 
leaves in a bud; 132, 133. 

Pubens, Pubes Used for Pubescent. 

Pub*rulus. Minutely pubescent. 

Pubes. Pubescence, hairiness. 

Pubescent (-ens). Clothed or furnished 
with hairs or down, especially with soft 
or downy and short hairs. 


Lying quite flat cn 


See Proter- 


Where flowering 


Alga, &c., 


GLOSSARY. 


Pugioniform (-ormis). Dagger-shaped. 

Pullus. Dark-colored; dusky-brown or 
blackish. 

Pulvereus, Pulverulentus. Powdered; as 
if dusted with powdery matter or 
minute grains. 

Pilvinate (-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. 

Pimilus. Low or little. 

Punetate (-atus). Dotted, either with 
depressions like punctures, or trans- 
lucent internal glands, or with colored 
(lots. 

Puneticulate (-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. 

Purpurascens. Purplish. 

Pusillus. Very small, or weak and slen- 
der. 

Pustular, Pustulate (-atus), 
(-osus). 
blisters. 

Putdmen. The shell of a nut; the endo- 
carp of a stone-fruit; 288. 

Pyenos, Greek for thick; whence Pyenv- 
cephalus, thick-headed, &c. 

Pygmeus. Dwarf, pygmy. 

Pyramidal (-alis). Pyramid-shaped. 

Pyréne (Pyréna). Same as Nucule or 
Nutlet; one of the small stones of a 
drupaceous fruit: 298. 

Pyrenarium, Pyridium. A pear or pear- 
like fruit, same as Pomum. 

Pyrenarvius. Name of a drupaceous 
pome, as of Medlar and Crategus. 

Pyridion. Synonym of Pome. 

Pyrenocarp (-arpium). A general name 
for any drupaceous fruit; 292. 

Pyriform (-ormis). See Pear-shaped. 

Pyzxidate (-atus). YVurnished with a lid. 

Pyzidium, Pyxis. A capsule with trans- 
verse dehiscence, making a lid of the 
upper portion; 293. 


Pustulose 
Having low elevations, like 


Quadri-. In Latin compounds, denotes 
four; as Quadrangular, Quadrifiri- 
ous (in four vertical ranks), Qudriju- 
gate (in four pairs), &c. 


429 


Quaternary, Quaternate. In fours or 
composed of four; 176. 

Quini, Quinary (-ius), Quinate (-atus). 
In fives; 176. 

Quinqgue. Five. In Latin compounds, 
giving rise to such terms as 

Quincuncial, in a Quincunz; also five- 
ranked; 125, 156. 

Quinquefarious (-ius), 
ranks. 

Quinquefoliate (-atus). Five-leaved. 

Quinquefoliolate, with five leaflets. 

Quintuple. Dividing into five parts, or 
five-fold. 

Quintuplinerved or -veined. With mid- 
rib of leaf dividing into five (i.e. two 
lateral pairs) above the base; 93. 


In five vertical 


Race. A variety of such fixity that it 
is reproduced by seed; also used ina 
looser and more extended sense for a 
series of related individuals without 
particular regard to rank; 320. 

Raceme (Racemus). An indeterminate 
or centripetal form of inflorescence 
with lengthened axis and equal-pedi- 
celled flowers; 146. 


| Racemiferous. Bearing racemes. 


Racemiform (-ormis). In the form of a 
raceme. 

Racemose (-osus). Having the character 
or appearance of a raceme, or in ra- 


ceines. 


Rachis. See Rhachis 
Radial. Belonging to the ray. 


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 eapitu- 
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 (-alis). 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. 

Radiciflorous (-us). Flowering (appar- 
ently) from the root. 

Radiciform (-ormis), Radicinus. Of the 
nature or appearance of a root. 

Radicle (4eula). Literally a diminutive 
root; but the “‘radicle’’ of the embryo, 


450 


so called in descriptive botany, is the 
hypocotyledonary and primal inter- 
node. (See Caulicle); 10. 

Radiculose (-osus). Bearing rootlets. 

Rudiz. The root. 

Rameal (-alis), Rameus. 
(Ramus) a branch. 

Ramenta. Thin chaffy scales belonging 
to the surface or epidermis, such as 
the chaff on the stalks of many Ferns. 

Ramification. Branching; 47. 

Ramijlorous (-us). Flowering on the 
branches. 

Ramose (-osus). Branching or branchy. 

Ramulose (-vsus). Bearing many branch- 
lets, ¢. e. Ramuli or 

Riphe. See Rhaphe. 

Rephides or Rhaphides. Crystals in 
ihe cells of plants, especially needle- 
shaped crystals. 

Ray (Radius). One of the radiating 
branches of an umbel (147); also the 
niarginal 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 

Ray-flowers. Those which belong to 
the margin of a circular flower-clus- 
ter, and differ from (being usually 
larger than) those of the disk. 

Recaulescence. The adhesion of leaves 
or their stalks to a stem; 158. 

Receptacle (Recepiaculum). 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 the 
axis or rhachis of the head, spike. or 
other dense cluster; 143. 

Reclinute (-atus\, Reclined, Reclining. 
Falling or turned toward downward, 
so that its upper part rests on the 
ground or other object; 53, 133. 

Rectinervius. Straight-veined or straight- 
nerved; 92. 

Rectiserial (-ialis). In rectilinear ranks ; 
124. 

Recurved (-us), Recurvatus. 
backward or downward. 

Reduplicate (-atus) or Reduplicativus. 
Folded and projecting outward. 

Reflexed (-us). Abruptly bent or turned 
downward or backward. 

R. fracted (-us). Same as reflexed, but 
abruptly bent from the base. 


Belonging to 


Curved 


GLOSSARY. 


Reygm1. A two-several-lobed two- 
several-celled fruit (2-pluricoccous), 
which separates at maturity into as 
many 2-valved carpels, as in Euphor- 
bia; one form of Schizocarp. 

Regmacarp (-arpium). A general name 
of a dry and dehiscent fruit, 292. 

Regular (-aris). Uniform in shape or 
structure; symmetrical as respects 
shape; 175. 

Reniform (-ormis). Kidney-shaped; 
having the outline of the longitudinai 
section of a kidney; 96. 

Repand ( Repandus). With slightly un- 
even margin, which, if more pro- 
nounced, would be sinuate; 98. 

Repent (Lepens). Creeping, 7. €. pros- 
trate or horizonal and rooting; 53. 

Replicate (-atus), Replicativus. Yolded 
backwar.!. 

Replum. <A frame-like placenta (like 
a door-case), from which the valves 
of a capsule or other dehiscent fruit 
fall away in dehiscence, as in Cruci- 
fere, certain Papaveraceez, Mimosa, 
&c.; 293. 


Reptant (Reptans). Same as Repent. 


Resupinate. Upside down, or having 
that appearance. 
Rete. Network. 


Reticulated (-atus), Retiformis. 
form of network; netted. 

Reticulate-veined, 92. 

Retindculum. Name sometimes applied 
to the gland to which one or more 
pollinia are attached in Orchids, &e. 
The persistent and indurated hook- 
like funiculus of the seeds in most 
Acanthacee. 

Retinerved (-ius). 
veined. 

Retrocurved (-us). Same as Recurved. 

Retroflexed (-us). Same as Reflexed. 

Retrorse (-orsus). Directed backward or 
downward. 

Retroverted (Retroversus). Inverted. 

Retuse (Retusus). With a shallow or 
obscure notch at a rounded apex; 97. 

Reversion A changing back, or in the 
reverse direction: 171. 

Rerolute (-utus). Rolled backward from 
the margins or apex; 133. 

Rhachis. The axis (backbone) of a 
spike or of a compound leaf; 101, 143. 

Rhaphe. The adnate cord or ridge 
which in an anatropous ovule con- 
nects the hilum with the chalaza; 
279, 307. 

Rhipidium. A fan-shaped cyme; 156. 


In the 


Same as Reticulate- 


GLOSSARY. 43 


Rhizanthous (-us). Root-flowered ; flower- 
ing from the root or seeming root. 

Lhizina. 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, Rhizima. A _ rootstock; a 
stem of root-like appearance pros- 
trate ou or underground, from which 
rootlets are seut off; the apex pro- 
gressively sending up herbaceous 
stems or flowering stalks and often 
leaves; 56. 7 


Rhizomorphous (-us). Root-like in ap- 


pearance. 
Rhombic (-icus). Rhomb-shaped. 
Rhomboidal (-alis). 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. 

Rimuse (-osus). With chinks or cracks, 
like those of old bark. 

Ring. In Ferns, &c. See Annulus. 

Ringent (-ens). Grinning or gaping ; as 
is the mouth of an open bilabiate 
corolla; 248. 

Riparius. Growing along the banks of 
rivers, &c. 

Rirdlis Growing along brooks. 

Rirvularis. 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 
mostly developed underground, and 
absorb moisture, &c., from the soil; 
2G 

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 rosé; 246. 
Sometimes used for rose-color. 


Roseus. Rose-colored; pale red. 

Rostellate (-atus). Diminutive of Ros- 
trate. 

Rostéllum. A diminutive beak. Also 


the name applied by Linneus to the 
Caulicle or Radicle. 

Rostrate (-itus). With a Rostrum, a 
beak or spur; narrowed into a slender 
tip or process. 

Rosular, Rosulate (-atus). 
a rosette. 

Rotate (-atus). Wheel-shaped; circular 
and horizontally spreading very flat ; 
248. 

Rotund (Rotundus, Rotundatus). Round- 
ed in outline: 95. 

Rough, Roughish, See Scabrous. 

Rubellus, Rubescent (-ens), Rubens. Red- 


Collected in 


dish. Rubescent also is turning red. 
Ruber. Red in general. 
Rubicundus Blushing, turning rosy- 
red. 


Rubiginose (osus). Brownish rusty-red. 

Ruderal (-alis). Growing in waste places 
or among rubbish. 

Rudiment. An imperfectly developed 
and functionally useless organ; a 
Vestige. 

Rufous (-us), Rufescent (-ens). 
red mixed with brown. 

Rugose (-osus). Covered or thrown into 
wrinkles, Ruge. 

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. 


Pale 


Ruptilis. Bursting irregularly. 
Rusty. Same as Rubiginose, Rufescent, 


and Ferruginous. 


Rutiluns. Deep red with a metallic 
lustre. 
Sabulosus. Growing in sandy places. 


Saccate (-atus), Sacciform. Sac-shaped ; 
baggy. 

Sagittate (-atus), Sagittiform (-ormis). 
Arrow-head-shaped. 

Salsuginosus. Growing within reach of 
salt water. 
Salver-shaped. 

phous; 248. 


See Hypocraterimor- 


452 


Samara. 
294. 

Samaroid. Resembling a samara. 

Sap-wood. New wood of an exogenous 
stem; 80. 

Sarcocarp (-arpium). The succulent or 
fleshy portion of a drupe; 285. Has 

een proposed also as a general name 
for a baccate fruit; 292. 

Sarmentose’ (-osus). Producing long 
and lithe branches or runners, viz. 
Sarments (Sarmenta). 

Sativus. That which is sown or planted. 

Saw-toothed. See Serrate. 

Saxdatilis, Suxosus, Saxicolus. Living on 
or among rocks. 

Scabridus, Scabriusculus. 
diminutive of 

Seabrous (Scaber). Rough to the touch. 

Scalariform (-ormis). Ladder-shaped; 
with transverse markings like the 
rounds of a ladder. 

Scales. Any thin scarious bodies, usu- 
ally degenerate leaves, sometinves of 
epidermal origin. 

Scalloped. Same as Crenate; 98. 

Scaly. See Scarious, Squamoese. 

Scaly Buds, 40. 

Scandent (-ens). 
ever mode; 51. 

Scape (Scapus). A peduncle rising from 
the ground; 51, 143. 

Scapiform (-ormis), Scapose (-osus). Re- 
sembling a scape. 

Scapigerous (-us). 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 Scariose (-osus). Thin, dry, 
membranaceous, and not green. 

Schizocarp (-arpium). <A pericarp which 
splits into one-seeded pieces; 296. 

Scion. A young shoot; a twig used for 
grafting. 

Sciuroideus. Like a squirrel’s tail. 

Scleranthium. Name of the fruit of 
Mirabilis, and the like; an akene 
enclosed in an indurated portion of 
calyx-tube. 

Scleroideus. Having a hard texture; 
from Scleros, hard. 

Scobiform (-ormis). Having the appear- 
ance of sawdust. 

Seorpioid. A form of unilateral inflo- 
rescence which is cireinately coiled in 
the bud: in the stricter sense, a form 
with the flowers two-ranked, these 
being thrown alternately to the right 
and left; 155, 157. 


An indehiscént winged fruit; 


Roughish; 


Climbing, in what- 


GLOSSARY. 


Scrobiculate ( atus). Marked by minute 
or shallow depressions. 

Scrotifurm (-ormis). Pouch-shaped. 

Scurf. Small and bran-like scales on 
the epidermis. 

Scutate (-atus), ~Scutiform (-ormis). 
Buckler-shaped. 

Scutelliform (-ormis). Platter-shaped. 

Scymetar-shaped. See Acinaciform. 

Sectile (-ilis). As if cut up into portions. 

Section (Sectio). In classification, is 
applied in a general way to a divi- 
sion in the arrangement of genera, 
species, or other groups; 327. 

Sectus. Completely divided; 99. 
Secund (Secundus). 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 
phenogamous plait; 305. 

Seed-leaves. Cotyledons, 11. 


Seed-stalk. See Funiculus and Podo- 
sperm. 

Seed-vessel. See Pericarp. 

Seyetalis. Growing in grain-fields. 

Segment (Segmentum). One of the 


divisions into which a plane organ, 
such as a leaf, may be cleft. 


Segregate (-atus). Separated; kept 
apart. 
Semen. Seed. 


Semi. Half, in Latin compounds: such as 

Semi-adherent. The lower half adhe- 
rent, &e.; Semi-amplezicaul (-aulis), 
half clasping the stem ; Semiovate, 
ovate halved lengthwise, &c. 

Semiandtropous. Same as Amphitro- 
pous: 279. 

Semilunar, Semilunate (-atus). A syn- 
onym of Lunate, being like a half- 
moon. 

Seminal (-alis). 

Seminiferous (-us). 

Sempervirent (Sempervirens). 
green. 

Senary (-arius). 

Sepal ( Sepalum). 

Sepaline (-inus), Scpalous. 
sepals. 

Sepaloid (-oideus). Resembling a sepal. 

Sepalody. Name for the metamorphosis 
of petals, &e., into sepals or sepaloid 
organs; 174. : 

Separated flowers. Those of distinct 
sexes; same as Diclinous; 191. 


Relating to the seed. 
Seed-bearing. - 
Ever- 


In sixes; 176. 
A calyx-leaf; 165. 
Relating to 


GLOSSARY. 


Septate (-atus). 
tion or septum. 

Septicide, Septiculal (-cidus). When a 
capsule dehisces through the dissepi- 
ments or lines of junction ; 289. 

Septiferous (-us). Bearing the partition 
or dissepiment. 

Septifragal (-us). Where the valves in 
dehiscence break away from the dis- 
sepiments ; 290. 

Septum. Any kind of partition, whether 
a proper dissepiment or not, 

Septulate (-atus). Divided by spurious 
or transverse septa. 

Serial (Serialis) or Seriate (Seriatus). 
Disposed in series or rows, whether 
transverse or longitudinal. 

Sericeous (-cus). Silky; clothed with 
close-pressed soft and straight pubes- 
cence. 

Serotinous (-us). Produced compara- 
tively late in the season. 

Serr:te (-atus). Beset with antrorse 
teeth; 97. 

Serrulate (-atus). Serrate with 
small or fine teeth; 97. 

Sesqui. <A Latin prefix denoting one 
and a half; as, Sesquipedalis, a foot 
and a half. 

Sessile (-ilis). Sitting close, without a 
stalk ; destitute of peduncle, pedicel, 
or petiole, as the case may be. 

Sets. A bristle, or bristle-shaped body. 

Sctaceous (-eus). Bristle-like. 

Setiform (-ormis). In the form of a 
bristle. 

Setigerous (-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, Sexpartile, &c. 

Shaggy. Pubescent with long and soft 
hairs; same as Villous. 

Sheath. 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. Having the character of a shrub. 

S/eve-cells, 77. 


Separated by a parti- 


very 


435 


Sigillate (-alus). 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. 

Sil culusa. Name of the Linnean arti- 
ficial order of the class Tetradynannia, 


having Siliculose pods ; 337. 
Silique (Siliqua). The peculiar pod of 
Crucifere, especially when mech 


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, 
&e. <A simple pistil is of one carpel; 
a simple leaf, of one blade, &e. 

Simple Fruits, 291. 

Simplicissimus. Most simple; complete- 
ly simple. 

Sinistrérse. Turned or directed to the 
left; 51, 140. 

Sinuate (-atus). 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. €. not rough, or to glabrous, 7. e. 
not pubescent ; the former is the more 
correct application. 

Soboles. Shoots, especially those from 
the ground. 

Soboliferous (-us). 
lithe shoots. 

Solid Bulb. A corm; 61. 

Solitary (-arius). Single, only one from 
the same place. 


Bearing vigorous 


Solubilis. Separating into portions or 
pieces. 
Solitus. Loosed; becoming separate. 


Sordidus. Of a dull or dirty hue. 

Sorcdiate (-atus). Bearing small patches 
on the surface. 

Sorema. A heap of carpels belonging to 
one flower; 2635. 

Sori, sing. sorus. Heaps, such as the 
clustered fruit-dots of Ferns. 

Sorose. Heaped or bearing Sori. 

Sorésis. A fleshy multiple fruit, such 
as a mulberry, bread-fruit, and pine- 
apple. 

Spadiceus. A bright and clear brown, 
or chestnut color. 


454 


Spadiceous. 
bearing a 

Spadic. A spike with a fleshy axis; 149. 

Span. The length of the space between 
the tip of the thumb and that of the 
little tinger, when outstretched ; about 
nine inches. 

Sparsus. Sparse or scattered; whence 
Sparsiflorus, with scattered flowers; 
Sparsifvlius, with scattered leaves, &c. 

Spathaceous (-us). Spathe-bearing, or 
of the nature of a 

Spathe (Spatha). A large bract, or a 
pair of bracts, enclosing a flower-clus- 
ter; 142. 

Spathella. An unused name for the 
glumes of Grasses. 

Spathilla. 
spathe. 

Spatulate (Spathulatus). Oblong with 
the lower end attenuated, shaped like 
a druggist’s spatula; 96. 

Species. The particular kind, the unit 
in natural history classification; 317. 

Specific Character, Name, &c., 349, 
363. 

Spermaphore or Spermophore (-orum). 
A name for the Placenta. 

Spermoderm (-ermis). The outer seed- 
coat; 305. 

Spermodophorum or Sperméphorum. An 
unused name for the gynophore in 
Umbellifere. The latter also an un- 
used name for the Placenta; 261. 

Spermothéca. An unused name for peri- 
carp. 

Spermum. Latin form of the Greek word 
for seed. Lat. Semen. 

Sphalerocarpium. Name proposed for 
an accessory fruit, such as that of 
Shepherdia, in which an akene is 
enclosed in a baccate calyx-tube. 

Spica. See Spike. 

Spicate (-atus). In the form of or resem- 
bling a spike, or disposed in spikes. 

Spiciform (-ormis). Spike-like. 

Spicula. .A diminutive or secondary 
spike; a Spikelet. 

Spike (Spica). A form of indeterminate 
inflorescence, with flowers sessile on 
an elongated common axis; 149. 

Spikelet (Spicula). A secondary spike; 
the name given to the Locusta or clus- 
ter of one or more flowers of Grasses 
subtended by a common pair of 
glumes. 

Spindle-shaped. See Fusiform. 

Spine (Spina). A sharp-pointed woody 
or indurated body,commonly a branch, 


Having the nature of or 


A secondary or diminutive 


GLOSSARY. 


sometimes a petiole, stipule, or other 
part of a leaf; 55, 117. 

Spinescent (-ens). Ending in a spine 
or sharp point; 59. 

Spinose (-osus). Furnished with spines, 
or of a spiny character; 55. 

Spinuliferous or Spinulose (-osus). Fur- 
nished with diminutive spines or Spin- 
ule. 

Spiral (Spirdlis). As if wound round 
an axis. Spiral Ducts, 68. Spiral 
Phyllotaxy, 119, 121. 

Spiricles. The delicate coiled threads 
in the hairs on the surface of certain 
seeds and akenes, which uncoil when 
wet; 307. 

Spithameus. A span long; the length 
spanned between the tip of thumb and 
forefinger when extended. 

Splendens. Resplendent or glittering. 

Spongelet, Spongiole (-icla). 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). 
analogue of seed in Cryptogams. 

Spore-case. See Sporangium. 

Sporidium. Synonym or diminutive of 
Spore. 

Sporiferous. Spore-bearing. 

Spérocarp (-arpium). Name given to 
certain spore-cases, as of Lycopodi- 
ace. 

Sporophore (-orum). 
nyms of Placenta. 

Sporule (Spdérula). Diminutive spore or 
a sort of spore. 

Sporuliferous (-us). 
taining spores. 

Sport. A bud-variation or seed-varia- 
tion; 319. 

Spumescent (-ens), Spumose. 
in appearance. 

Spur. <A hollow and slender extension 
of some portion of the blossom, usu- 
ally nectariferous, as of the calyx of 
Larkspur and the corolla of Violet: 
rarely applied also to a solid spur-like 


The 


One of the syno- 


Bearing or con- 


}Froth-like 


process. 

Spurred. Producing a spur. See Cal- 
carate. 

Squama. <A scale of any sort, usually 


the homologue of a leaf. 
Squamate (-atus), Squamiferous, Squa- 
mosus. Furnished with scales. . 


aeale 


GLOSSARY. 425 
Squamella, Squamul. Diminutive compounds as Distichous, two-ranked ; 


squama; scales of secondary order 
or reduced size. 

Squamiform (-ormis). 

Squamulose (-osus). 
with minute scales. 

Squarrose (-osus). Literally rough- 
scurfy; applied to bodies rough with 
spreading and projecting processes, 
such as tips of bracts, Ke. 

Squarrulose (-osus). Diminutively squar- 
Tose. 

Stachys. Greek for spike. 

Stalk. Any kind of lengthened support 
on which an organ is elevated. 

Stamen. One of the elements or phylla 
of the andreccium ; 165. 

Staumineal, Stamineous (-eus). 
to the stamens; 151. 

Staminifcrous (-us). Stamen-bearing. 

Staminodium. 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. 


Seale-like. 
Covered or beset 


Relating 


Stans. Supporting itself in an erect 
position. 
Station. Particular place as to soil, ex- 


posure, &e., which a plant affects; 


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. 

Stenos. Greek for narrow; hence 

Stenophyllus. Narrow-leaved, &c. 

Sterigma. Any foliaceous prolongation of 
the blade of a leaf down on the stem 
by decurrence. 

Stériymum. Name of Desyaux for the 
Dieresilis of Mirbel. 

Sterile (-ilis). Barren. as a blossom 
destitute of pistil, 191; a stamen 
without anther, or an anther without 
pollen; an ovary, without good ovules, 
seeds without embryo, &e. In com- 
mon English use. a male or staminate 
‘lower is said to be a sterile flower. 

Stichvs. Greek for row or rank, usually 
meaning vertical rank ; hence such 


Tristichous, three-ranked, &c. 

Stigma, pl stigmata. 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. 

Stigmatic (-icus), Stigmatose (-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 Pern ; any 
stalk-like support of a gyncecium or 
a carpel; 212. 

Stipel (Stipelium). An appendage to a 
leaflet analogous to the stipule of a 
leaf; 106. 

Stipellate (-atus). Provided with stipels ; 
106. 

Stipitate (-atus). 
special stalk. 
Stipiiifjorm (-ormis). Shaped like a stipe; 

stalk-like. 

Stipulaceous (-eus), Stipular (-aris). 
Belonging to stipules. 


Having a stipe or 


Stipulate. Possessing stipules. 
Stipules. Appendages or adjuncts of a 


leaf one ou each side of the insertion ; 
85, 105. 

Stirps, pl. 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. 

Stoloniferous (-us). Sending of or propa- 
gating by stolons, runners, &e. 

Stoma, pl. stémata, 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 


Strap-shaped. See Ligulate (247) and 
Lorate. 

Striate (-atus). Marked with fine longi- 
tudinal lines, streaks, or diminutive 
grooves or ridges (Strie). 

Strict (Strictus). Close-or narrow and 
upright; very straight. 

Striyillose (-osus). Minutely strigose. 

Strigose (-osus). Beset with strige, or 
sharp-pointed and appressed straight 
and stiff hairs or bristles. 

Strobilaceous (-cus), Strobiliform (-or- 
mis). Relating to or 1esembling a 
Strobile. 

Strobile (Stidbilus). An inflorescence 
formed largely of imbricated scales, 
as that of Hop and a Fir-cone; 
303. 

Strombuliformis, Strombuliferus. Twist- 
ed spirally into a serew shape, as the 
legumes of the Screw-bem (Proso- 
pis, sect. Strombocarpa) and of some 
species of Medicago. 

Strdéphiole (-iola). 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 struma or goitre-like 
swelling. 

Stupose (-osus). Tow-like; with tufts or 
mats of long hairs. 

Style (Stylus). The usually attenuated 
portion of a pistil or carpel between 
the ovary and stigma; 166. 

Styliform (-ormis). Style-shaped. 

Styliferous. Style-beariug. 

Stylinus. Belonging to the style. 

Stylosus. With styles of remarkable 
length or number, &e. 

Stylopodium. Anenlargement ora disk- 
like expansion at the base of a style, 
as in Umbellifere. 

Sub. In composition of Latin words in 
terminology, denotes somewhat or 
slightly; as, Subacute, Subcordate, that 
is acutish, somewhat cordate, Ke. 

Subclass, 327. 

Subconvolute and Subimbricate in eestiva- 
tion, 137. 

Suberose (-osus). Of a corky texture. 

Subgenus, 327. 

Submerged, Submersed (-us). 
ufder water. 

Suborder (Subordo), 327. 

Subpetiolar (-aris). Under the petiole, 
as the leaf-buds of Platanus; 42. 

Subsection, 327. 


Growing 


| 
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} 


GLOSSARY. 


Subspecies. A group which is ambigu- 
ous in rank between variety and spe- 
cies; 320. 

Subtribe (Subtribus), 327. 

Subulate (-atus), Subuliform (-ormis). 
Awl-shaped. 

Subvariety, 527. 

Succise (-isus). As if eut or broken off 
at the lower end. 

Succubous (-us). When in_ leaves 
crowded on a stem the apex of each 
leaf is covered by the base of the 
next above. 

Succulent (Succosus). Juiey. 

Sucker. A shoot of subterranean ori- 
gin; 53. 

Suffrutescent (-ens). 
scurely shrubby; 50. 

Suffrutez. An undershrub. 

Suffruticose (-osus). Low and shrubby 
at base; 50. 

Suffultus. _Underpropped or supported. 

Sulcate (-atus). 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; 160, 183. 

Superposed (Superpositus). 
over some other part. 

Superposition, 179, 195. 

Supervolute (-us), Supervolutive (-ivus). 
Same as Convolute when applied to 
plaits; 139. 7 

Supine (-inus). Lying flat with face up- 
ward. 

Suppression. Complete abortion ;~179, 
190. 

Supra. Above; hence in Latin com- 
pounds, Supra-axillary, above the 
axil; Suprafoliaceous,above a leaf, &e. 

Supradedecompound. Several times com- 
pound. 

Surculose (-osus). Producing suckers. 

Surculus. A sucker; a shoot rising from 
a subterranean base; 53. 

Sursum. Upward; directed upward or 
forward. 

Suspended (Suspensus). Hanging di- 
rectly downward; hanging from the 
apex of a cell. 

Suspensor of the embrvo, 284. 

Sutural (-alis). Relating to a suttire. 

Suture (-ura). A junction or seam of 
union; used commonly as a line of 
opening; 260. e 

Sword-shaped. A blade with two sharp 
and neaily parallel edges, as in Iris. 


Slightly or ob- 


Vertically 


GLOSSARY. 


Syconium or Syconus. A multiple fruit 
like that of the Fig; 148, 303. : 

Sylvestrts. 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 orgaus 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. With stamens united. 

Sympode, Sympodium. A stem made 
up of a series of superposed branches 
in a way 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 6n the angles. 

Synema. The column of monadelphous 
filaments, as in Mallow. 

Syngenesia. Linnzan class (335) charac- 
terized by having the anthers united or 

Syngenesious. With anthers cohering 
in aring; 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). 
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. Sameas Fulyous; dull brown- 
ish-vellow. 


Wasting or shrivel- 


= Tendril. 


| 


437 


Taxology, Taxonomy. Relating to clas- 
sification and its rules; 3, 315. 

Teeth. Any small marginal lobes. 

Teymen. The inner coat of a seed; 306. 

Tela. Latin name for tissue, cellular 
tissue, Ke. 

Teleianthus. Same as perfect, or her- 
maphrodite-flowered. 

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). 

Teratclogical. 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. 

Fernate (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 
Tetracarpellary (-aris). 
pels; 261. 


Of four car- 


Tetracamarous (-us), Tetracoccus. Of 
four closed carpels. 
Tetradyndmia. Linnean class (335) 


which has the stamens. 
Tetradynamous (-us). With four long 
and two shorter stamens; 250. 


Tetragonal or Tetragunous (-us). Four- 
angled. 
Tetragynia. Linnean artificial order 


(337), characterized by having the 
gynecium. 
Tetrdgynous. Of four carpels or styles. 
Tetramerous (-us). Composed of four 
members in a circle; 176. 


Tetrandria. Linnean class having the 
flowers perfect and 

Tetrandrous. With four stamens; 249, 
334. 


438 


Tetrapetalous (-us). 
244, 

Tetraphyllous (-us). 

Tetraquctrous (-us). 
or salient angles. 

Tetrasepalous (-us). 
244. 

Tetrdstichous (-us). In 
ranks. 

Thalamiflorous (-us), 340. 
of the flower hypogynous, or on the 
Thalamus. The receptacle of a flower; 

167. See Torus. 

Thallophytes ( Thallophyta), 341. 

Thallus. A stratum, in place of stem 
and foliage. 

Theca. Acase: an anther-cell (251); 
a spore-case, &c. (An early name for 
the anther, 166.) 

Thecaphore (-orum). 


With four petals; 


Four-leaved; 245. 
With four sharp 


With four sepals; 


four vertical 


The stipe of a 


earpel (homologous with petiole); 
212. 
Thorn. Same as spine; 55. 


Throat. ‘The orifice of a gamopetalous 
corolla or calyx, including any por- 
tion between this and the proper tube; 
246. See Faux. : 

Thyrse, Thyrsus. A contracted or ovate 
panicle; a mixed inflorescence, with 
inain axis indeterminate, but the sec- 
ondary or ultimate clusters cymose ; 
15). 

Tigelle, Tigellula. A miniature or ini- 
tial stem ; sometimes applied to Cauli- 
cle (Radicle), sometimes to Plumule; 
10. 

Tinctorius. Dyed; used for dyeing; 
imparting color. 

Tissue. The anatomical fabric. 

Tomeéntose (-osus). Densely pubescent 
with matted wool, or Tomentum. 

Tongue-shaped. Long and nearly flat, 
somewhat fleshy, and rounded at the 
apex. 

Tooth. See Teeth. 

Tvothed. See Deutate. 

Top-shaped. Junversely conical. 

Torose (-osus). Cylindrical, with con- 
tractions or bulges at intervals. 

Tortuous (-osus). Bent or twisted in 
different directions. 

Torulose  (-osus). 
slightly torose. 


Diminutively or 


Tortus. Twisted. 

Tortilis. Susceptible of twisting. 

Torus. The receptacle of a flower; 167, 
211. 


Trabeculate (-atus). Cross-barred. 
Trachea. Aspiral vessel or duct, named 


With parts 


GLOSSARY. 


from resemblance to the trachee of 
insects. 

Trackycarpous (-us) Rough-fruited. 

Trachyspermous. Rough-seeded, &e. 

Transverse (-ersus). Across; right and 
left as to bract and axis; collateral; 
160. 

Trapeziform (-ormis), Trapezoid. Un- 
symmetrically four-sided, like a tra- 
pezium. 

Tree. A woody plant with an elevated 
trunk. 

Tri-. In compound words, both Latin 
and Greek, denotes three or triple. 

Triachanium. A fruit like a cremocarp 
but of three carpels. 

Triadelphous, Triadelphia. 
iments in three sets; 250. 

Triandria. Linnean class (3384) with the 
flowers. 

Triandrous. With three stamens; 249. 

Triangular (-aris), Triangulatus. Three- 
angled. 

Trianthous (-us). Three-flowered. 

Tribe. Group superior to genus, in+ 
ferior to order; 326. 

Tricarpellary (-aris). 
261. 

Tricarpous (-us). 
fruits or carpels. 

Tricephalous (-us). Bearing three heads. 

Trichocarpous (-us). Hairy-fruited. 

Trichodes. Resembling hair. 

Trichdtomous (-us). Three-forked ; 
branched into three divisions. 

Tiichome (Trichéma). Any outgrowth 
of the epidermis, such as a hair or 
bristle; 209. 

Tricoccous (-us). 
cocci. 

Tricolor. Three-colored. 

Tricuspidate (-atus). Tipped with three 
cusps or pointed tips. 

Tridéntate (-atus). Three-toothed. 

Tridigitate. Thrice digitate. 

Triduus. Lasting for three days. 

Tricnnial (Triennis). Lasting for three 
years. 

Tiifarious (-ius). Facing three ways, 
in three vertical ranks. 

Trifid (Ti ifidus). Three-cleft. 

Trifoliate (-ius, Trifoliatus). 
leaved. 

Trifoliolate (-atus). 

Trifiircate (-atus). 
forks or branches. 

Trigamous (-us). Bearing three kinds 
of flowers. 

Tiigonous (-us), Trigonal. Three-angled. 


With fila- 


Of three carpels; 


Consisting of three 


Consisting of three 


Three- 


Of three leaflets. 
Divided into three 


GLOSSARY. 


Trigynia. Linnean artificial order with 
Trigynous, zi. e. three-styled flowers; 
337. 

Trihilatus. Waving three apertures, as 
in some grains of pollen. 

Trijugate (Trijugus). With three pairs 
of leaflets or pinuz. 

Trilobate (Trilubus). Three-lobed. 

Trilocular (-aris). Three-celled. 

Trimerous (-us). Three-membered parts 
in threes; 176. 

Trimestris. Lasting for or maturing in 
three months. 

Trimorphous, Trimorpism. 
under three forms; 236. 

Trinervate (Trinervius). 

Trinodal. Of three nodes or joints. 

Triecia. Linnean artificial order with 
the flowers. 

Triecious or Trivicous (-us). Having 
staminate, pistillate and perfect flowers 
(or three kinds of flowers as to sex), 
334; on three distinct plants. 

Tridvulate (-atus). Having three ovules. 

Tripartible (-ibilis). Tending to split 
into three portions. 

Tripartite (-itus). ‘Three-parted. 

Tripetalvid (-videus). As if three-pet- 
alled. 

Tv ipetulous(-us). Having three petals. 

Triphyllous (-us). Three-leaved; 243. 

Tripinnate (-atus). Thrice pinnate; 104. 

Tripinnatifid (-idus). Thrice pinnatifid. 

Triple-ribbed or nerved. With midrib 
dividing into three, or sending off on 
each side a strong branch, above the 
base of the blade; 93. 

Triplinerved (Triplinervius). Same as 
Triple-nerved, Triple-ribbed; 93. 

Tripterous (-us). Three-winged. 

Triquetrous (Triqueter). Three-edged ; 
with three salient angles. 

Triquinate (-atus}. Divided first into 
three then into five. 

Trisected (-us). Divided into three por- 
tions; 99. 

Trizepalous (-us). Of three sepals. 

Triserial (-alis), Triseriate (-atus). In 
three horizontal ranks or series. 

Tristachyus. Three-spiked. 

Tristichous (-us). In three vertical 
ranks; 122. 

Tristigmatic. With three stigmas. 

Tristis. Dull colored. 

Tristylous (-us). Having three styles. 

Trisilcate (-atus). Three-grooved. 

Triternate (atus). Thrice ternate; 104. 

Trivial names, Nomina trivialia. Com- 
mon or yulgar names; used by Lin- 


Occurring 


Three-nerved. 


439 


neus for specific names of a single 
word; 346, 362. 

Trochlear (-earis). Pulley-shaped. 

Trophosperm (Trophospermium). Name 
for the Placenta; 261. 

Trumpet-shaped. ‘Tubular, with a dilat- 
ed orifice. 

Truncate (-atus). 
end; 97. 

Trunk (Truncus). A main stem. 

Tryma. A drupaceous nut, with exo- 
carp at length dehiscent or otherwise 
separating, such as walnut and hick- 
ory nut. 

Tubeformis. Trumpet-shaped. 

Tube (Tubus). Any hollow elongated 
body or part of an organ; 245. 

Tuber. A thickened and short subter- 
ranean branch, beset with buds or 
eyes; 59. 

Tubercle (Tuberculum). A small tuber 
or an excrescence: or something be- 
tween a tuber and a root; 60. 

Tuberculate (-atus). Beset with knobby 
projections or excrescences. 

Tuberiferous. Bearing tubers. 

Tubular, Tubulosus (-ose). Having a 
tube; tube-shaped; 248. 

Tubuliflorus (-us). When the flowers of 
a head have only tubular corollas. 

Tunicate (-us). Having coats (tunics). 

Turbinate (-atus). Top-shaped. 

Turion, (Turio). A scaly sucker or 
shoot from the ground; 41. 

Turnip-shaped. See Napiform. 


As if cut off at the 


Twin. In pairs. See Geminate, Didy- 
mous. 
Twining. Winding -spirally and so 


climbing (Twiners); 51. 
Twisted. Contorted. 
Two-lipped. See Bilabiate. 
Type. The ideal plan or pattern. 
Typical. Representing the plan or 
type. 


Uliginose (-osus). Growing in swamps. 

Ulnaris. Of the length of the ulna or 
fore-arm. 

Umbel (Umbélla). An inflorescence 
(properly of the indeterminate type) 
in which a cluster of pedicels spring 
all from the same point, like rays of 
an umbrella; 146. 

Umbellate (-atus). Umbellitorm (-ormis). 
In or like umbels. 

Umbellet. A partial or secondary um- 
bel; 150. 


Umbellifercus (-us). Bearing umbels. 


440 GLOSSARY. 


Umbéllula. A partial or secondary um- 
bel, or umbellet; 150. 

Umbilicate (-atus). Depressed in the 
centre, navel like. 

Umbilicus. The hilum of a seed. 

Umbonate (-atus). Bearing an Umbo 
or loss in the centre 

Umbraculiform (-ormis). Having the 
general form of an umbrella. , 

Umbrosus. Growing in shady places. 

Unarmed. Destitute of prickles, spines, 
or other armature. 

Uncate (-atus), Uncinate, (-atus), Unei- 
form (-ormis). Hooked; bent or 
curved at tip in the form of a hook. 

Uncialis. An inch (uncia) in length. 

Undute (-atus) or Undulate (-tus). 
Wavy; 98. 

Undershrub. A very low shrub; 50. 

Unequally pinnate. See Impari-pin- 
nate. 

Unguiculate (-atus). Contracted at 
base into an 

Unguis. A claw, or stalk-like base of a 
petal, &c.; 245. 

Uni-. In Latin compound, one; as 

Unicellular. Of one cell; Unicolor, of 
one color, &e. 

Unicus. Singly or single, solitary. 

Uniflorous (-us). One-flowered. 

Onifoliate (-atus). One-leaved. 

Unifoliclate, of one leaflet; 102. 

Unijigate (Unijugus). Of one pair; 
102. 

Unilabiate (-atus}. One-lipped, like the 
corolla of Acanthus, in which the 
upper lip is obsolete. 

Unilateral (-alis). One-sided; either 
originating on or more commonly 
turned all to one side of an axis. 

Onilocular {-aris). One-celled. 

Uninervate (Uninervis, Uninervius). 
One-nerved. 

Uniovulate (-atus). Having only a soli- 
tary ovule. = 

Uniparous. Bearing one; as a cyme of 
one axis or branch; 152, 155. 

Uniserial (-ialis), Uniseriate (-atus). 
In one horizontal row or series. 

Unisexual (-alis, Uniserus). Of one 
sex; haying stamens only or pistils 
only; 191. 

Univalved ( Univalvis). Of one piece or 
valve. 

Ureevlate (-atus) Hollow and con- 
tracted at or below the mouth, like an 
urn or pitcher ( Urceolus). 

Urens. Stinging, in the manner of net- 
tles. 


Utricle (Utriculus}. A small bladdery 
pericarp; 295. Orany small bladder- 
shaped body or appendage; also a 
synonym of a cell of parenchyma. 

Utricular (-aris), Utrieulate (-atus), 
Utriculiform (-ormis), Utriculose 
(-osus). Havirg or consisting of 
utricles, or bladder-like in appear- 
ance. 


Vacillans. Swinging free, as the anth- 
ers of Grasses on their filaments. 

Vacuus. Void or empty of the proper 
contents. 

Vagina. A sheath, as of a leaf, &e. 

Vaginate. Sheathed. 

Vall cule. The intervals or grooves 
between the ridges or ribs of the fruit 
Umnbellifere. 

Valcate (-atus), Valvular (-aris). 
Opening as if by doors or valves, as 


do most dehiscent fruits (capsules), 


and some authers; also the parts of 
a flower-bud when they exactly meet 
without overlapping; 135. 

Valve (Valva). One of the pieces into 
which a capsule splits, 288. 

Valved. Same as valvate: hence 3 
-valved, 5-valyed, many valved, &e. 
Valeula. A diminutive valve. Also 
used (after Linnzus) for the inner or 

flower-glumes of Grasses. 

Variegated (-atus). Irregularly colored ; 
in patches of color. 

Variety (Varietas). A sort or modifi- 
cation subordinate to species; 318. 

Varivlate, Variolaris, Marked as if 
by the pustules or pittings of small- 
pox. 

Vascular (-aris). Relating to or fur- 
nished with vessels ( Vasa) or ducts. 

Vascular Plants ( Vasculares), 340. 

Vasculum. Same as Ascidium. Also 
the botanists’ collecting box ; 372. 

Vasiform (-ormis). In the form of a 
vessel, duct, &e. 

Veined. Furnished or traversed with 
fibro-vascular bundles or threads, es- 
pecially with those which divide and 
are reticulated. 

Veins (Vene). In general any ramifi- 
cations or threads of fibro-vascular 
tissue in a leaf or any flat organ; 
especially (as distinguished from 
nerves) those which divide or branch ; 
92. 

Veinless. Destitute of veins. 

“einlet (Venula). Gne of the ultimate 


ow fe — wren tet pn” * * 


1 cnt er INN aoa = 


on 


GLOSSARY. 


or smaller ramifications of a vein or 
rib; 93. 

Velute (-atus). Veiled. 

Veliutinous (Velutinus). Velvety: the 
surface covered with a soft coating of 
fine and close silky pubescence, or 
velumen. 

_ Venation ( Venatio). 
ing; 90. 

Venenatus, Venenosus. Poisonous. 

Venose (-osus) Veiny; abounding in 
veins or network. 

Ventral (-alis). Belonging to the an- 
terior or inner face of a carpel, Kc.; 
the opposite of dorsal. 

Ventricose (-osus). Swelling unequally 
or inflated on one side. 

Ventriculose (-osus). Minutely ventri- 
cose. : 

Venulose (-osus). Abounding with vein- 
lets or venule. 

Vermicular (-aris) | Worm-shaped. 

Vernal (Vernalis), Appearing in spring. 

Vernation (-ativ). The disposition of 
parts in a leaf-bud; 132. 

Vernicose (-vosus). As if varnished. 

Veérrucose (-osus). Covered with warts 
(verruce) or wart-like elevations. 

Versatile (Versatilis). Swinging toand 
fro; turning freely on its support; 
253. 

Versicolor. Changing color, or of more 
than one tint cr color. 

Vertex. The apex of au organ. 

Vertical (-alis). Perpendicular to the 
horizon; longitudinal. 

Verticil (-illus). A whorl; 6. 

Verticillaster. A false whorl, composed 
of a pair of opposite cymes; 159. 

Verticillastrate. Bearing or arranged 
in Verticillasters. 

Verticillate (-atus, -aris). 
whorl; 6, 119, 120. 

Vescicle (-icula). A small bladder or 
air-cavity. 

Vesicular (-aris), Vesiculose (-osus). As 
if composed of little bladders. 

Vespertine (Vespertinus). Appearing or 
expanding in early evening. 

Vessels (Vase). See Ducts. 

Verillary (-aris), Vesxillar, 137. Per- 
taining to the 

Vezillum. The standard or large pos- 
terior petal of a papilionaceous corolla; 
184. © 

Villose (osus) or Villous. Bearing shaggy 
or long and soft (not interwoven) hairs 
or Villi. 

Vimineous(-eus). Bearing long and flex- 


The mode of vein- 


Disposed in a 


441 


ible twigs, like those used for wicker 
work. : 

Vine. Any trailing or climbing stem: 
originally that of the Grape from 
which wine is made. 

Vinealis. Growing in vineyards. 

Violaceous (-eus). Violet-colored. 

Virens. Green, or evergreen. 

Virescens. Greenish or turning green. 

Virgate (-atus). Wand-shaped, or like 
a rod; slender, straight, and erect. 

Virgultum. A vigorous twig or shoot. 

Viridescent (-ens). Same as Virescens. 

Viridis. Green. 

Viridulus. Greenish. 

Virosus. Yenomous. 

Viscid (-idus), Viscous (-osus). Sticky 
from a tenacious coating or secretion. 


Vitellinus. The yellow hue of the yolk 
of egg. 
Vitéllus. Name formerly given to the 


peculiar albumen which is in some 
cases deposited within the embryo-sac. 

Viticulose (-osus). Sarmentaceous ; pro- 
ducing vine-like twigs or suckers, 
viticule. 

Vitte. The fillets or stripes (vil-tubes) 
of the pericarp of most Umbelliferz, 
which contain an aromatic or peculiar 
secretion. 

Vittate (-atus). Bearing vittee; or with 
any longitudinal stripes. 

Viviparous (-us). Germinating or sprout- 
ing from seed or bud while on the 
parent plant. 

Volutle ( Volibilis). 
support; 51. 

Volutus. Rolled wp in any way. 

Volva. A wrapper or external covering, 
especially that of many Fungi. 


Twining round a 


Wavy. See Undulate. 

Waszy. Resembling beeswax in consist- 
ence or appearance. 

Wedge-shaped or Wedge-form. See 
Cuneate; 95. 

Wheel-shaped. See Rotate. 

Whorl. Arranged in a circle round an 
axis; a Verticil; 6. 

Whorled. Disposed in whorls. 


Wild. Growing without cultivation; 
spontaneous. 
Winy. See Ala. Any Membraneous 


or thin expansion by which an organ 
is bordered, surrounded, or otherwise 
augmented. Also the two lateral 
petals of a papilionaceous corolla are 
termed wings; 185. 


442 


Winged. See Alate: bearing a wing or 
wings. 

Withering. See Marcescent. 

Wood. The hard part of a stem, &c., 
mainly composed of 

Wovod-cells, Woody fibre or tissue, 68. 

Woolly. See Lanate and Tomentose: 
clothed with long and tortuous or 
matted hairs. 


Xanthos. Greek for yellow in com- 
pounds, such as Xanthephyll, the yel- 
low coloring matter in leaves. 


GLOSSARY. 


Xenogamy. Fecundation of the ovules of 
a flower by pollen from some other 
plant of the same species; cross-fer- 
tilization; 216. 

Xylinus. Woody, pertaining to wood. 


Zoospore. One of the free-moving spores 
of the lower Cryptogams. 

Zygomorphous (-us). That which can be 
bisected in only oue plane into similar 
halves; 175. 


ADDENDA. 


Antidromous, Antidromy. 
Infertile (-iiis). 


When the course of a spiral is reversed, 157. 
Said of a pistil or flower which fails to set fruit. 


Polyembryony. The production of two or more embryos in a seed, 284. 


Saprophytes (-yta). 


Plants feeding upon decaying vegetable or animal matter. 


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