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Professor of Botany in the University of Copenhagen 



M. C. POTTER, M.A. F.L.S. 

Professor of Botany in the University of Durham 

College of Science, Newcastle-upon-Tyne 
Author of "An Elementary Text-book of Agricultural Botany' 










THE present translation of Dr. E. Warming's Haandbog i den 
Systematiske Botanik is taken from the text of the 3rd Danish 
Edition (1892), and from Dr. Knoblauch's German Edition (1890), 
and the book has been further enriched by numerous additional 
notes which have been kindly sent to me by the author. Dr. 
Warming's work has long been recognised as an original and 
important contribution to Systematic Botanical Literature, and I 
have only to regret that the pressure of other scientific duties 
has delayed its presentation to English readers. Dr. Warming 
desires me to record his high appreciation of the careful transla- 
tion of Dr. Knoblauch, and his obligation to him for a number of 
corrections and improvements of which he has made use in the 3rd 
Danish Edition. In a few instances I have made slight additions 
to the text ; these, however, appear as footnotes, or are enclosed 
in square brackets. 

In the present Edition the Thallophytes have been revised and 
rearranged from notes supplied to me by Dr. Knoblauch, to whom 
I am indebted for the Classification of the Fungi, according to 
the more recent investigations of Brefeld. The Bacteria have 
been revised by Dr. Migula, the Florideae rearranged after Schmitz, 
and the Taphrinacese after Sadebeck. The main body of the text 
of the AlgfE and Fungi remains as it was originally written by 
Dr. Wille and Dr. Rostrup in the Danish Edition, though in many 
places considerable alterations and additions have been made. 
For the sake of comparison a tabular key to the Classification 
adopted in the Danish Edition is given in the Appendix. 

In the Angiosperms I have retained the sequence of orders in 

the Danish original, and have not rearranged them according to 
w. B. v I 


the systems more familiar to English students. In any rearrange- 
ment much of the significance of Dr. Warming's valuable and 
original observations would have been lost, and also from a 
teacher's point of view I have found this system of great value. 
Although at present it may not be completely satisfactory, yet as 
an attempt to explain the mutual relationships, development and 
retrogression of many of the orders, it may be considered to have 
a distinct advantage over the more artificial systems founded 
upon Jnssieu's Divisions of Polypetala?, Gamopetala?, and 

With reference to the principles of the systematic arrangement 
adopted, I may here insert the following brief communication 
from the author (dated March, 1890), which he has requested me 
to quote from the preface of Dr. Knoblauch's edition : " Each 
form which, on comparative morphological considerations, is 
clearly less simple, or can be shown to have arisen by reduction 
or through abortion of another type having the same fundamental 
structure, or in which a further differentiation and division of 
labour is found, will be regarded as younger, and as far as 
possible, and so far as other considerations will admit, will be 
reviewed later than the ' simpler,' more complete, or richer forms. 
For instance, to serve as an illustration : EPIGYNY and PEKIGYNY 
are less simple than HYFOGXY ; the Epigynous Sympetalie, Chori- 
petalte, Monoctyledones are, therefore, treated last, the Ilydro- 
charitacem are considered last under the Helobiet?,etc. ZYGOMOIM'HY 
is younger than ACTINOMORPHY ; the Scilaminece and Gynandrce 
therefore follow after the LiliijJora', the Scrophulariacere after the 
Solanacecc, Linaria after V&rbascum, etc. FORMS WITH UNITED LEAVES 
indicate younger types than those with free leaves ; hence the 
SympetalcB come after the Choripetaltv, the Silenece after the 
Alsinex, the Nal 'cacetv after the Stcrculiacex and Tiliacecc, etc. 

" ACYCLIC (spiral-leaved) flowers are older than cyclic (verticillate- 
leaved) with a definite number, comparing, of course, only those 
with the same fundamental structure. The Veronica-type must 
be considered as younger, for example, than Digitalis and Antir- 


rli-inum, these again as younger than Scrophularia Verbascum, on 
the contrary, is the least reduced, and therefore considered as the 
oldest form. Similarly the one-seeded, nut-fruited Ranunctdacew 
are considered as a later type (with evident abortion) than the 
many-seeded, folicular forms of the Order ; the Paronychiece and 
Chenopodiacece as reduced forms of the Alsinece type ; and the 
occurrence of few seeds in an ovary as generally arising through 
reduction of the many-seeded forms. The Cyperacece are regarded 
as a form derived from the Jimcacece through reduction, and 
associated with this, as is so often the case, there is a complication 
of the inflorescence ; the Dipsacacece are again regarded as a form 
proceeding from the Valerianactce by a similar reduction, and 
these in their turn as an off-shoot from the Caprifoliacece, etc. Of 
course these principles of systematic arrangement could only be 
applied very generally ; for teaching purposes they have often 
required modification." 

In preparing the translation considerable difficulty has been 
experienced in finding a satisfactory rendering of several terms 
which have no exact equivalent in English. I may here especially 
mention the term Vorblatt (Forblad) which I have translated by 
the term bracteole, when it clearly applied to the first leaf (or 
leaves) on a pedicel ; but in discussing questions of general 
morphology a term was much needed to include both vegetative 
and floral shoots, and for this I have employed the term " Fore- 
leaf." Also, the term " Floral-leaf " has been adopted as an 
equivalent of "Hochblatt," and the term "bract" has been limited 
to a leaf subtending a flower. I have followed Dr. E. L. Mark 
in translating the word " Anlage " by "Fundament." 

At the end of the book will be found a short appendix giving 
an outline of some of the earlier systems of Classification, with a 
more complete account of that of Hooker and Bentham. 

In a book of this character it is almost impossible to avoid 
some errors, but it is hoped that these will be comparatively few. 
In correcting the proof-sheets I have received invaluable assist- 
ance from Dr. Warming and Dr. Knoblauch, who have kindly 


read through every sheet, and to whom I am greatly indebted for 
many criticisms and suggestions. I have also to thank Mr. I. H. 
Burkill for his kind assistance in looking over the proofs of the 
Monocotyledons and Dicotyledons, and Mr. Harold Wager for 
kindly reading through the proofs of the Algce and Fungi. My 
thanks are also especially due to Mr. E. L. Danielsen, and I wish 
to take this opportunity of acknowledging the very considerable 
help which I have received from him in translating from the 
Original Danish. 

January, 1895. 



(The Alc/CE and Fungi re-arranged in co-operation with Dr. E. Knoblauch, the 
other Divisions as in the 3rd Danish Edition.) 


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

B. Sub-Division. Algae. ......... 8 

Class 1. SYNGENETKLE 14 


,, 3. DIATOMEJE 18 


Family 1. Schizophyceos 22 

2. Bacteria 26 

Class 5. CONJUGATE 41 


Family 1. Protocoecoidese 47 

,, 2. Confervoidese ....... 53 

3. Sipboneee 59 

Class 7. CHAEACE.E 64 


Family 1. Pba3osporea3 68 

,, 2. Cyclosporese 73 

Class 9. DICTYOTALES 76 


Family 1. Bangioidese 77 

,, 2. Florideffi 78 

C. Sub-Division. Fungi 84 


Sub-Class 1. Zygomycetes 96 

,, 2. Oomycctes 100 

Family 1. Eutoinophthorales 102 

2. Cbytridiales 102 

,, 3. Mycosiphonales 104 




Class 2. MESOMYCETES ... ... . 108 

Sub-Class 1. Hemittsci 108 

,, 2. Hemibasidii ..... . 109 

Class 3. MYCOJIYCETI s (HIGHER FUNGI) . .... 114 

Sub-Class 1. Ascvmycetes . 114 

Series 1. Exoasci ... . . 116 

2. Carpoasci ... .118 

Family 1. Gjumoascales 118 

,, 2. Perisporiales . .... 119 

,, 3. Pyrenomycetes 125 

,, 4. Hysteriales ....... 132 

,, 5. Discomycetes .... . 132 

6. Helvellales . . ... 136 

Ascolichenes 136 

Sub-Class 2. Basidiomycetes ... . 144 

Series 1. Protobasidoinycetes . . 145 

,, 2. Autobasidiomycetes . 157 

Family 1. Dacryomycetes . 159 

,, 2. Hynienoinycetes . . 159 

3. Phalloideffi . 172 

,, 4. Gasteromycetes . . 173 

Basidioliclienes . 176 

Fungi Imperfecti . .176 


Class 1. HEPATIOE 188 

Family 1. Marcbautiese ... . 190 

,, 2. Antbocerotese .191 

., 3. Jungermanmere ... 191 

Class 2. Musci FRONDOSI. .... . 192 

Family 1. Spbagnete . .193 

,, 2. Schizocarpeae . 

,, 3. Cleistocarpeas . . 

,, 4. Stegocarpete ... 195 


Class 1. FILICIN.E .... 205 

Sub-Class 1. Filices . . 205 

Family 1. Eusporangiatse. . . 210 

,, 2. Leptosporangiatfe . . 212 

Sub-Class 2. Hydroptendece . 215 
Sub Class 1. Isosporous Equiset line 

2. Heterosporous Equisttince . 225 

Class 3. LYCOPODIN.E (CLUB MOSSES) . . 226 

Sub- Class 1. Lycopodiece ... 226 

,, 2. Selayinellea 228 


Asexual Generation of the Cormopbytes . 
Sexual Generation ; Fertilisation .... 

Class 1. CYCADEJE (CYCADS) ...... 

2. CONIFER.E (PlXE-TREES) ..... 

Family 1. Taxoideas ...... 

,, 2. Piiiouleffi ...... 

Class 3. GXETE.E 

Fossil Gymnosperms ..... 



Family 1. Helobieas ...... 

,, 2. Glumifiora} ...... 

3. Spadicifloraa ..... 

4. Euautioblastas . . . 

,, 5. LiliifloriB ...... 

6. Scitamineas ..... 

,, 7. Gynandrae ...... 

ClaSS 2. DlCOTYLEDONES ...... 

Sub-Class 1. Choripctiiltc ...... 

Family 1. Saliciflorte ...... 

,, 2. Casuariniflorre ..... 

,, 3. Quercifloras ..... 

4. Juglandifloraj ..... 

,, 5. Urticiflorse . ..... 

0. Polygonifloraa ..... 

,, 7. Curvembryaj ..... 

,, 8. Cactiflorce 

,, 9. Polycarpicas ..... 

,, 10. Rhosadinaa ...... 

,, 11. Cistiflorag ...... 

12. Gruinales ...... 

,, 13. Columniferaj ..... 

14. Tricoccos 

,, 15. Terebinthiua) 

,, 16. Aesculinas ...... 

,, 17. Frangulinte 

,, 18. Tliymelasinas 

,, 19. Saxifragiua) ..... 

,, 20. Bosiflorse 

,, 21. Legumiuosaa ..... 

,, 22. Passiflorinas ..... 

,, 23. Myrtiflora3 ...... 

24. Umbellifloraa 

,. 25. Hysterophyta ..... 


. 234 

. 234 

. 243 

. 251 

. 252 

. 255 

. 259 

. 262 

. 270 

. 271 

. 273 

. 274 

. 278 

. 283 

. 297 

. 308 

. 309 

. 323 

. 328 

. 334 

. 337 

. 337 

. 339 

. 340 

. 349 

. 351 

. 358 

. 363 

. 375 

. 377 

. 393 

. 406 

. 416 

. 421 

. 430 

. 435 

. 439 

. 443 

. 448 

. 451 

. 456 

. 466 

. 475 

. 482 

. 490 



Sub-Class 2. Sympetalce 

A. Pentacyclicic . 
Family 26. Bicornes . 

,, 27. Diospyrina: 
,, 28. Primulinffl 

B. Tetracycllccc . 
Family 29. Tubiflorte 

,, 30. Personata? 
,, 31. Nuculifera3 

32. Contorts; . 

33. Eubiales . 
34. Dipsacales 

,, 35. Campannlinffi . 
36. Aggregates 



. 504 

. 506 

. 506 

. 510 

. 511 

. 514 

. 514 

. 517 

. 531 

. 541 

. 548 

. 556 

. 560 

. 564 

. 574 


Page 9, line 12 from top, for Hydrodicton read Hydrodiclyon. 

,, 14, lines 1 and 2 from top, for as in the preceding case read in tltis case. 

,, 14, ,, 2 and 15 from top, for zygote read oosporc. 

,, 88, line 15 from bottom, for Periphyses read periphyses. 

,, 124, ,, 7 ,, ,, for CJucromyces read Choiromyces. 

,,142, ,, 2 ,, and in Fig. 137, for Bnomyves read Bccomyccs. 

,, 152, ,, 2 ,, top, for Pints read Pi/nts. 

,, 152, ,, 5 ,, for Crategus read Cratagus. 

216, Fig. 215, for Salcina read Salvinia. 

,, 306, line 6 from top, for Pista read Pistia. 

,,316, ,, 26 ,, after Dracaena insert a comma. 

,, 337, ,, 13 ,, ,, for end read beginning. 

,, 483, ,, 11 ,, bottom, for Lagerstnvmia read Lagcrstrameria. 
For a, 6 and ii read as, ce and ue throughout. 

The following are not officinal in the British Pharmacopoeia : page 316, 
Dractcna (Dragon's-blood), Smilax glalra ; p. 321, " Orris-root " ; p. 326, species 
of Curcuma, Alpinia qfficinarum; p. 333, Orr/tj's-species ("Salep"). On page 
296, par. 4, only Pearl Barley is offic. iu the Brit. Phar. 


The Vegetable Kingdom is arranged in 5 Divisions. 

Division I. Thallophyta, Stemless Plants, or those which 
are composed of a " thallus," i.e. organs of nourishment which 
are not differentiated into root (in the sense in which this term is 
used among the higher plants), stem, or leaf. Vascular bundles 
are wanting. Conjugation and fertilisation in various ways; 
among most of the Fungi only vegetative multiplication. 

In contradistinction to the Thallopbytes all other plants are called " Stem- 
plants" (" Cormophyta "), because their shoots are leaf-bearing stem?. The 
name Thallophyta (Stemless-plants) is to some extent unsuitable, since many 
of the higher Alga? are differentiated into stem and leaf. 

The Thallophytes are again separated into 3 sub-divisions, 
namely : 

Sub-Division A. Myxomycetes, Slime-Fungi, with only 1 


Sub-Division B. Algse, with 10 classes : 
Class 1. Syngeneticaa. 

,, 2. Dinoflagellata, Peridinea. 
,, 3. Diatomese, Diatoms. 
,, 4. Schizophyta, Fission Algae. 
,, 5. Conjugate. 
,, 6. Chlorophycene, Green Algse. 
,, 7. Characeae, Stone-worts. 
,, 8. Phteophycea?, Brown Alga 1 . 
,, 9. Dictyotales. 
,, 10. Rhodophycere, Red Algaa. 
Sub-Division C. Fungi, with 3 classes: 
Class 1. Phycomycetes. 
,, 2. Mesomycetes. 
3. Mycomycetes, Higher Fungi. 

Division II. Bryophyta or Muscineae, Mosses. These 
have leaf-bearing shoots, but neither true roots nor vascular 
w. B. B 


bundles. The lowest Mosses have, however, a thallus. Fertili- 
sation is accomplished by means of self-motile, spirally coiled 
spermatozoids, through the agency of water. From the fertilised 
oosphere a " fruit-body " (capsule) with unicellular organs of 
reproduction (spores) is produced. The spore on germination 
gives rise to the vegetative system, which bears the organs of 
sexual reproduction ; and this system is divided into two stages 
the protonema, and the leaf-bearing plant produced on it. 
Alternation of generations : 

I. The protonema and the entire nutritive system which 

bears the organs of sexual reproduction. 
II. The capsule-like sporangium, with spores. 

2 Classes : 1. Hepatica?, Liverworts. 

2. Musci, Leafy Mosses. 

Division III. Pteridophyta or Vascular Cryptogams, 
Fern -like Plants having- leaf-bearing shoots, true roots, and 
vascular bundles with tracheides and sieve-tubes. Fertilisation 
as in the Mosses. From the fertilised oosphere the leaf -bearing 
shoot arises, which bears on its leaves the reproductive organs, 
the spores, in capsule-like sporangia. From the germination of 
the spore a small prothallium is formed, which bears the sexual 
reproductive organs. 

Alternation of generations : 

I. Prothallium with organs of sexual reproduction. 
II. Leaf-bearing shoot with capsule-like sporangia. 

3 Classes: 1. FilicinaB, True Ferns. 

2. Equisetina?, Horsetails. 

3. Lycopodinse, Club-mosses. 

Division IV. Gymnospermae. The vegetative organs are in 
the main similar to those in the 3rd Division ; special shoots are 
modified into flowers for the service of reproduction. From the 
oosphere, which is fertilised by means of the pollen-tube, the 
leaf -bearing plant is derived; this passes the first period of its 
life as an embryo in the seed, and continues its development when 
the germination of the seed takes place. The organs correspond- 
ing to the spores of the two preceding Divisions, are called respec- 
tively the pollen-grain and embryo-sac. The pollen-grains are 
multicellular ; i.e. they contain an indistinct prothallium. In the 
embryo-sac a prothallium, rich in reserve material (endosperm), 


with female organs of reproduction, is developed BEFORE FER- 
TILISATION. The pollen-grains are carried by means of the wind 
to the ovules; these enclose the embryo-sac, and are situated on 
the open fruit-leaf (carpel), which has no stigma. 
Alternation of generations : 

I. Prothallium = Endosperm in ovule. 

II. Leaf-bearing plant, with flowers which produce the pollen- 
sac and ovule. 
3 Classes : 1. Cycadeae. 

2. Coniferae. 

3. Gnetaceee. 

Division V. Angiospermae. The members of this group are 
very similar to those of Division IV. The ovules are, however, 
encased in closed fruit-leaves (ovary), which have a special portion 
(stigma) adapted for the reception and germination of the pollen- 
grains. The pollen-grains are bicellular, but with only a mem- 
brane separating the two nuclei ; they are carried to the stigma 
by animals (chiefly insects), by the wind, or by some other means. 
Endosperm is not formed till AFTER FERTILISATION. Alternation of 
generations in the main as in the Gymnosperms, but less distinct ; 
while the sexual generation, the prothallium, with the organs of 
fertilisation, is also strongly reduced. 

2 Classes: 1 1. Monocotyledones. Embryo with one seed-leaf. 
2. Dicotyledones. Embryo with two seed-leaves. 

For a long time the vegetable kingdom has been divided into CRYPTOGAMS (so 
called because their organs of reproduction remained for some time undis- 
covered), and PHANEROGAMS or Flowering-plants which have evident sexual 

The first three divisions belong to the Cryptogams, and the third and fourth 
divisions to the Phanerogams. This arrangement has no systematic value, but 
is very convenient in many ways. 

The Cryptogams are also known as Spore-plants, since they multiply by 
unicellular organs (spores), and the Phanerogams in contradistinction are 
called Seed-plants (Spermaphyta), since they multiply by seeds, multicellular 
bodies, the most important part of which is the embryo (a plant in its 
infancy). Mosses, Ferns, and Gymnosperms are together known as Arche- 
goniatre, fiuce they possess in common a female organ of distinct structure, 
the Archegonium. 

1 See Angiospernise. 



The thallus in the simplest forms is unicellular; in the majority, 
however, it is built up of many cells, which in a few instances 
are exactly similar ; but generally there is a division of labour, so 
that certain cells undertake certain functions and are constructed 
accordingly, while others have different work and corresponding 
structure. Vessels or similar high anatomical structures are 
seldom formed, and the markings on the cell-wall are with few 
exceptions very simple. The Myxomycetes occupy quite an 
isolated position ; their organs of nourishment are naked masses 
of protoplasm (plasmodia). 

As regards the external form, the thallus may be entirely 
without special prominences (such as branches, members), but 
when such are present they are all essentially alike in their 
origin and growth, that is, disregarding the hair- structures 
which may be developed. A shoot of a Seaweed or of a Lichen, 
etc., is essentially the same as any other part of the plant ; only 
among the highest Alga? (Characete, certain Siphonese, Sar- 
gaxsum, and certain Red Seaweeds) do we find the same differ- 
ences between the various external organs of the plant body as 
between stem and leaf, so that they must be distinguished by 
these names. 

Roots of the same structure and development as in the Seed- 
plants are not found, but organs of attachment (rhizoids and 
haptera) serve partly the biological functions of the root. 

belong three sub-divisions Slime-Fungi, Alga?, and Fungi. 
Formerly the Thallophytes were divided into Alga?, Fungi, and 
Lichens. Biit this last group must be placed among the Fungi, 
since they are really Fungi, which live symbiotically with Alga 3 . 
The Slime-Fiingi must be separated from the true Fungi as a 
distinct sub-division. The Algce possess a colouring substance, 
which is generally green, brown, or red, and by means of which 
they are able to build up organic compounds from carbonic acid 
and water. The Bacteria, especially, form an exception to the 
Alga? in this respect ; like the Fungi and Slime-Fungi they have 
as a rule no such colouring material, but must have organic car- 
bonaceous food ; these plants form no starch, and need no light 


for their vegetation (most Fungi require light for fructification). 
The Myxomycetes, Bacteria, and Fungi derive their nourish- 
ment either as saprophytes from dead animal or vegetable matter, 
or as parasites from living animals or plants (hosts), in which they 
very often cause disease. 

A remark, however, must be made with regard to this division. Among the 
higher plants so much stress is not laid upon the biological relations as to divide 
them into " green " and " non-green"; Cuscuta (Dodder), a parasite, is placed 
among the Convolvulaceje, Neottia and Corallorhiza, saprophytes, belong to the 
Orchidacea 3 , although they live like Fungi, yet their relations live as Alga?. In 
the same manner there are some colourless parasitic or saprophytic forms among 
the Alga?, and stress must be laid upon the fact that not only the Blue-green Alg.'v, 
but also the Bacteria, which cannot assimilate carbonic-acid, belong to tlie Alga? 
group, Schizophycese. The reason for this is that systematic classifications must 
be based upon the relationship of form, development, and reproduction, and from 
this point of view we must regard the Bacteria as being the nearer relatives of 
the Blue-green Alga?. All the Thallophytes, which are designated Fungi (when 
the entire group of Slime-Fungi is left out), form in some measure a connected 
series of development which only in the lower forms (Phycomycetes) is related 
to the Algre, and probably through them has taken its origin from the Alga? ; the 
higher Fungi have then developed independently from this beginning. The 
distinction of colour referred to is therefore not the only one which separates the 
Algne from the Fungi, but it is almost the only characteristic mark by which 
we can at once distinguish the two great sub- divisions of the Thallophytes. 

The first forms of life on earth were probably " Protista?," which had assimi- 
lating colour material, or in other words, they were Algre because they could 
as-imilate purely inorganic food substances, and there are some among these 
which belong to the simplest forms of all plants. Fungi and Slime-Fungi must 
have appeared later, because they are dependent on other plants which assimi- 
late carbon. 1 


The Slime-Fungi occupy quite an isolated position in the 
Vegetable Kingdom, and are perhaps the most nearly related to 
the group of Rhizopods in the Animal Kingdom. They live in and 
on organic remains, especially rotten wood or leaves, etc., on the 
surface of which their sporangia may be found. 

They are organisms without chlorophyll, and in their vegetative 
condition are masses of protoplasm without cell-wall (plasmodt'a). 
They multiply by means of spores, which in the true Slime-Fungi 3 

1 According to the recent investigations of Winogradsky some micro-organisms 
(Nitrifying-bacteria) can build organic from inorganic matter. Sachs' hypothesis 
that the first organisms must necessarily have contained chlorophyll is there- 
fore untenable. 

2 Myxogasteres, Engler's Syllabus, p. 1. 



are produced in sporangia, but in some others * free. The spores 
are round cells (Fig. 1 a) which in all the true Slime-Fungi are 
surrounded by a cell-wall. The wall bursts on germination, and 
the contents float out in the water which is necessary for 
germination. They move about with swimming and hopping 
motions like swarmspores (e, /), having a cilia at the front end and 
provided with a cell-nucleus and a pulsating vacuole. Later on 

FIG. 1. o-l Development of " Fuligo " from spore to Mj-xamoeba ; a-m are magnified 
390 times ; m is a Myxamoeba, of Ltjcogala epidendron ; I' three Myxamoabfe of Physarum 
album about to unite; o, a small portion of plasmodiuni, magnified 90 times. 

they become a little less active, and creep about more slowly, while 
they continue to alter their form, shooting out arms in various 
places and drawing them in again (g, h, i, k, I, m) ; in this stage 
they are called Myxamcebce. 

The Myxamceba grows whilst taking up nourishment from the 
material in which it lives, and multiplies by division. At a later 
stage a larger or smaller number of Myxamcebae may be seen to 
1 Acrasiete aud Plasrnodiopkorales, Hid. 


coalesce and form large masses of protoplasm, plasmodia, which 
in the "Flowers of Tan" may attain the size of the palm of a 
hand, or even larger, but in most others are smaller. The plas- 
modia are independent, cream-like masses of protoplasm, often 
containing grains of carbonate of lime and colouring matter (the 
latter yellow in the Flowers of Tan). 
They creep about in the decaying matter 
in which they live, by means of amoeboid 
movements, internal streamings of the pro- 

FIG. 3. Pour sporangia of 
Stenwnitis fuscn, fixed on a 
branch, a The plasmodium. 

FIG. 2. The plasmodiinn fa) of Stemonitis fusea, com- 
mencing to form into sporangia (b) ; drawn on. July 9. 
The dark-brown sporangia were completely formed by 
the next morning ; e-e shows the development of their 
external form. 

toplasm continually taking place ; finally 
they creep out to the surface, and very 
often attach themselves to other objects, 
such as Mosses, and form sporangia (Fig. 
2). These are stalked or sessile and are 
generally cylindrical (Fig. 3), spherical 
or pear-shaped (Fig. 4); they rarely attain 
a larger size than that of a pin's head, and 
are red, brown, white, blue, yellow, etc., with a very delicate wall. 
In some genera maybe found a " Capillitium " (Fig. 4 cp), or net- 
work of branched fine strands between the spores. Flowers of 

FIG. 4. Sporangium of Ar- 
cyria incarnata. B closed; 
open ; p wall of sporangium ; 
cp capilitium. 



Tan (Fuligo septicd) has a fruit-body composed of many sporangia 
(an yEthalium), which has the appearance of flat, irregular, brown 
cakes, inside the fragile external layer of which a loose powder, 
the spores, is found. It generally occurs on heaps of tanners' bark, 
and appears sometimes in hot-beds in which that material is used, 
and is destructive by spreading itself over the young plants and 
choking them. 

All the motile stages may pass into resting stages, the small forms 
only surrounding themselves with a wall, but the large ones at the 
same time divide in addition into polyhedral cells. When favour- 
able conditions arise, the walls dissolve and the whole appears 
again as a naked (free-moving) mass of protoplasm. 

To the genuine Slime-Fungi belong : Arcyria, Trichia, Didymium, 
Physarum, Stemonitis, Lycogala, Fuligo, Spumaria, Reticularia. 

Some genera wanting a sporangium- wall belong to the Slime- 
Fungi : Ceratiomyxa, whose fruit-body consists of polygonal 
plates, each bearing stalked spores ; Dictyostelium, in which the 
swarm-stage is wanting and which has stalked spores. Plas- 
modiophora brassicce preys upon the roots of cabbages and other 
cruciferous plants, causing- large swellings. PL alni causes 
coral-shaped outgrowths on the roots of the Alder (Altius). 
Phytomysea Jeguminosariim may be found in small knobs (tubercles) 
on the roots of leguminous plants. It is still uncertain whether 
it is this Fungus or Bacteria which is the cause of the formation 
of these tubercles. 

Sub-Division II. ALG/E. 

Mode of Life. The Algte (except most of the Bacteria) are 
themselves able to form their organic material by the splitting 
up of the carbonic acid contained in the water, or air in some 
cases, and for this purpose need light. The majority live in 
water, fresh or salt, but many are present on damp soil, stones, 
bark of trees, etc. 

With the exception of the Bacteria, no saprophytes have actually 
been determined to belong to this group, and only very few 
true parasites (for instance, Phyllosiphon arisari, Mycoidea, etc.), 
but a good many are found epiphytic or endophytic on other Alga?, 
or water plants, and on animals (for instance, certain Schizophycece 
and Protocnccoidece ; Trichopliilus ivelckeri in the hairs of Bradypus, 
the Sloth), and several species in symbiotic relation to various 

ALG;E. 9 

Fungi (species of Lichen), to Sponges (e.g. Trentepohlia spongio- 
phila, Struvea delicatula), and to sundry Infusoria and other 
lower animals as Radiolarias, Hydra, etc. (the so-called Zoochlorella 
and Zooxantella, which are perhaps partly stages in development 
of various Green and Brown Alga>). 

Vegetative Organs. The cells in all the Alga? (excepting 
certain reproductive cells) are surrounded by a membrane which 
(with the exception of the Bacteria) consists of pure or altered 
cellulose, sometimes forming a gelatinous covering, at other times 
a harder one, with deposits of chalk or silica formed in it. The 
cell-nucleus, which in the Schizophyta is less differentiated, may 
be one or more (e.g. Hydrodicton, Siphonece) in each cell. Except- 
ing in the majority of the Bacteria, colour materials (of which 
chlorophyll, or modifications of it, always seems to be found) occur, 
which either permeate the whole cytoplasm surrounding the cell- 
nucleus, as in most of the coloured Schizophyta, or are con- 
tained in certain specially formed small portions of protoplasm, 

The individual at a certain stage of development consists nearly 
always of only one cell ; by its division multicellular individuals 
may arise, or, if the daughter-cells separate immediately after the 
division, as in many of the simplest forms, the individual will, 
during the whole course of its existence, consist of only a single 
cell (unicellular Alga?). In multicellular individuals the cells 
may be more or less firmly connected, and all the cells of the 
individual may be exactly alike, or a division of labour may take 
place, so that certain cells undertake certain functions, and are 
constructed accordingly ; this may also occur in parts of the cell 
in the large unicellular and multinuclear Alga? (Siphonea?, p. 62). 

The cells in most of the Alga? belong to the parencliymatous form ; 
these, however, in the course of their growth, may very often become 
somewhat oblong; in many Alga? (particularly Fucoidea? and 
Floridea>) occur, moreover, hyphce-like threads, which are very long, 
often branched, and are either formed of a single cell, or, more 
frequently, of a row of cells, having a well-pronounced apical 
growth. The parenchymatous as well as the hyphae-like cells 
may, in the higher Alga? (especially in certain Fucoidea? and 
Floridea?), be further differentiated, so that they form well- 
defined anatomico-physiological systems of tissue, i.e. assimilating, 
conducting, storing, and mechanical. 

With regard to the external form, the thallus may present no 



differentiation, as in many unicellular Alga 3 , or in multicellular 
Alga? of the lower order, which are then either equally developed 
in all directions (e.g. Pleurococcuf, Fig. 47), or form flat cell-plates 
(Merismopedium) or threads (Oscillaria, Fig. 21). The first step in 
the way of differentiation appears as a difference between apex 
and base (Sivularia, Porphyra) ; but the division of labour may 
proceed so that differences may arise between vegetative and 
reproductive cells (CEdogonium, Fig. 54) ; hairs and organs of 
attachment (rhizoids and haptera), which biologically serve as 
roots, are developed, and even leaves in certain forms of high 
order, belonging to different classes (e.g. Caulerpa, Fig. 59 ; 
Characeoe, Fig. 61 ; Sargassum, Fig. 72 ; and many Floridea?). 

The nonsexual reproduction takes place vegeiatively, in 
many instances, simply by division into two, and more or less com- 
plete separation of the divisional products (Diatomaceae, Desmi- 
diaceoe (Fig. 36), many Fission-plants, etc.), or by detached portions 
of the thallus (e.g. Caulerpa, Ulra lacttica, etc. ; among many Schizo- 
phyceSB, small filaments known as hormogonia are set free), or 
asexually by special reproductive cells (spores) set free from the 
thallus ; these may be either stationary or motile. The stationary 
reproductive cells (spores) may either be devoid of cell-wall (te- 
traspores of the Florideie), or may possess a cell-wall ; in the latter 
case they may be formed directly from the vegetative cells, gene- 
rally by the thickening of the walls (akinetes), or only after a 
process of re-juvenescence (aplanospores) . Aplanospores, as well 
as akinetes, may either germinate immediately or may become 
resting-cells, which germinate only after a period of rest. 

pear-shaped, naked, swarmspores (zoospores), which have arisen in 
other cells (zoosporangia), and propel themselves through the 
water by means of cilia ; or they are Phyto-Amceba?, which have no 
cilia and ci'eep on a substratum by means of pseudopodia. The 
cilia, which are formed from the protoplasm (in the Bacteria, 
however, from the membi'ane), are mostly situated at the pointed 
and colourless end, which is directed forwards when in motion, 
and are 1, 2 (Fig. 5 B), 4 or more. Both the cilia in the Brown 
Alga? are attached to one side (Fig. 65); they are occasionally 
situated in a circle round the fi'ont end (GEdogonium, Fig. 6 a, 
and Derbesia), or are very numerous and situated in pairs dis- 
tributed over a large part or nearly the whole of the zoospore 
(Vauclieria). Besides being provided with one or more nuclei 


(Vaucheria), they may also have a red "eye spot" and vacu- 
oles, which are sometimes pulsating, i.e. they appear and re- 
appear at certain intervals. The swarmspores move about in 
the water in irregular paths, and apparently quite voluntarily, 
revolving round their longer axes ; but they come to the sur- 
face of the water in great numbers either because of their 
dependence on light, or driven by warm currents in the 
water, or attracted by some passing mass of food material. 
The swarmspores germinate, each forming a ue\v plant, as their 
movement ceases they surround themselves with a cell-wall, 
grow, and then divide ; in Fig. 6 6, two may be seen in the con- 
dition of germination, and about to attach themselves by means 
of the front end, which has been developed into haptera (see also 
Fig. 5 B, lowest figure). 

FIG. 5. Cludopliora glomerata. A The lower cells are 
full of swarmspores, whilst from the upper one the 
greater part have escaped through the aperture m. 
B Free and germinating swarmspores. 

FIG. 6. (Edoqonium : a 
(free), b germinating swarm- 

The sexual reproduction here, probably in all cases, con- 
sists in the coalescence of two masses of protoplasm, that is, in 
the fusion of their nuclei. 

The simplest and lowest form is termed conjugation, orisoga- 
mous fertilisation, and is characterized by the fact that the two 
coalescing cells (termed gametes) are equal, or almost equal, in shape 
and size (the female gamete in the Cutleriacece, e.<j. Zanardinia 



collaris, Fig. 7, is considei^ably larger than the male gamete). The 
cell in which the gametes are developed is called a gametangimn, 
and the reproductive cell formed by their union which generally 

has a thick wall and only 
germinates after a short 
period of rest is termed a 
zyyote or sygospore. The con- 
jugation takes place in two 
ways : 

(a) In the one way the 
gametes are motile cells 
( planogametes, zoogametes, 
Fig. 8), which unite in 
pairs during their swarming 
hither and thither in the 
water; during this process 
they lie side by side (Fig. 8tZ), 
generally at first touching at 
the clear anterior end, and 
after a time they coalesce 
and become a motionless zy- 
fjote, which surrounds itself 
with a cell-wall (Fig. 8 e). 
This form of conjugation is found in Vlothrix (Fig. 8 d), Acetabu- 
laria, and other Alga? (Figs. 45, 56, 66). 

(b) Among other Alga? (e.g. Diatomacece and Conjugates), the 
conjugating cells continue to be surrounded by the cell-wall of 
the mother-cell (aplanogametes in an aplanogametanyinm] ; the 


FIG. 7. Zannnlinia co'ilaris. A Male game- 
tangia (the small-celled) and female gametangia 
(large-celled). C Female gamete. I) Male gamete. 
B E Fertilisation. F Zygote. G Germinating 


FIB. 8. UlotUrlx zoiinta: a portion of a thread with zoospores, of which two are formed 
in each cell (zoosporangium), the dark spots upon them are the " red eye-spots " ; 1, 2, 3, 
4 depict successive stages in the development of the zoospores ; b a single zoospore, at u 
the pulsating vacuole ; c portion of a thread with gametes, of which sixteen are formed in 
each g-ametangium ; d gametes free and in conjugation; e conjugation has been eSTected, 
and the formed zygotes are in the resting condition. 



aplanogametangia generally grow out into short branches, which 
lie close together and touch one another, the wall at the point 
of contact is then dissolved (Fig. 39). Through the aperture thus 
formed, the aplanogametes unite, as in the first instance, and 
form a rounded zygote, which immediately surrounds itself with 
a cell- wall. Vai-ious modifications occur ; compare Figs. 37, 39, 
41, 43. 

The highest form of the sexual reproduction is the Egg- or 
Oogamous fertilisation. The two coalescing cells are in the main 
unlike each other in form as well as size. The one which is con- 
sidered as the male, and is known as the spermatozoid (antherozoid), 
developes as a rule in large numbers in each mother-cell (antheri- 


FIG. 9. Fertilisation in the Bladder- 
wrack (Fucus vesiculosus). 

dinm) ; they are often self- 
motile (except in the Flori- 
deas, where they are named 
spermatici) , and are many 
times smaller than the other 
kind, the female, which is 
known as the eg g - cell 
(oosphere). The egg-cell is 
always a motionless, spheri- 
cal, primordial cell which can either float about freely in the 
water, as in the Fucaceje (Fig. 9), or is surrounded by a cell-wall 
(oogonium) ; generally only one oosphere is to be found in each 
oogonium, but several occur in Sphceroplea (Fig. 10). The re- 

FIG. 10. Spltosroplea annulina. 

14 ALG.E. 

suit of the spermatozoid coalescing with the egg-cell is, as in 
the preceding case, the formation of a zygote, which generally 
undergoes a period of rest before germination (the Florideae are 
an exception, a fruit-body, cystocarp, being produced as the result 
of coalescence). 

An example of fertilisation is afforded by the Alga, S^)llc^roplea anmdina (Fig. 
10). The filamentous thallus is formed of cylindrical ce Is with many vacuoles (r 
in A) ; some cells develope egg-cells (B), others spermatozoids (C), the latter in 
a particularly large number. The egg-cells are spherical, the spermatozoids of 
a club- or elongated pear-shape with two cilia at the front end (G ; E is however 
a swarmspore). The spermatozoids escape from their cells through apertures in 
the wall (o in C) and enter through similar apertures (o in .B) to the egg-cells. 
The colourless front end of the spermatozoid is united at first with the "re- 
ceptive spot" of the egg-cell (see f), and afterwards completely coalesces with 
it. The result is the formation of a zygote with wart-like excrescences (Z>). 

The female (parthenogenesis) or male (androgenesis) sexual cell 
may, sometimes without any preceding fertilisation, form a new 
individual (e.g. Ulothrix zonata, Cyhndrocapsa, etc.). 

Systematic division of the Algae. The Algae are divided 
into the following ten classes : 


Among the lowest forms of the Algae, the Syngeneticaa, the 
Dinoflagellata, and the unicellular Volvocaceae (Chlamydomonea?), 
distinct transitional forms are found approaching the animal 
kingdom, which can be grouped as animals or plants according to 
their method of taking food or other characteristics. Only an 
artificial boundary can therefore be drawn between the animal 
and vegetable kingdoms. In the following pages only those forms 
which possess chromatophores, and have no month, will be con- 
sidered as Alga?. 

Class 1. Syngeneticae. 

The individuals are uni- or multi-cellular, free-swimming or 
motionless. The cells (which in the multicellular forms are loosely 
connected together, of ten only by mucilaginous envelopes) are naked 
or surrounded by a mucilaginous cell- wall, in which silica is never 
embedded. They contain one cell-nucleus, one or more pulsating 




PIG. 11. Syncrypta volvox : the multi- 
cellular individual is surrounded by a 
mucilaginous granular envelope. 

vacuoles, and one to two band- or plate-like chrornatophores with 
a brown or yellow colour, and sometimes a pjrenoid. 

Reproduction takes place by vegetative division, or asexually by 
zoospores, akinetes (or aplanospores ?). Sexual reproduction is un- 
known. They are all fresh water forms. 

To this class may perhaps be assigned the recently arranged and very 
little known orders of Calcocytacece, Murracytacece, Xanthellacece, and 
Dictyochacece, which partly occur in the 
free condition in the sea, in the so-called 
" Plankton," and partly symbiotic in 
various lower marine animals. 

The Syngeneiicce are closely re- 
lated to certain forms in the animal 
kingdom, as the Flagellatse. 

Order 1. Chrysomonadinaceae. In- 
dividuals, uni- or multi-cellular, swim- 
ming in free condition, naked or sur- 
rounded by a mucilaginous covering. The 
cells are generally oval or elongated, with 
2 (rarely only 1) cilia, almost of the same 
length, and generally with a red " eye-spot " at their base, and with 2 (rarely 1 
only) band-shaped chromatophores. Eeproduction by the longitudinal divi- 
sion of the individual cells either during the swarming, or during a resting 
stage ; in the multicellular forms also by the liberation of one or more cells, 
which in the latter case are connected together. 

A. Unicellular: Chromulina, Cryptogleiia, Microglena, Nephroselmis. 

B. Multicellular: Uroglena, Syncrypta (Fig. 11), Synnra. 

Among the unicellular Chrysomouadinaceae are probably classed some forms 
which are only stages in the development of the multicellular, or of other 

Order 2. Chrysopyxaceae are unicellular, and differ mainly from the pre- 
ceding in being attached either on a slime-thread (titylo- 
chry sails), or enclosed in an envelope (Chrt/sopyxis, Fig. 
12). They have two cilia, and multiply by longitudinal 
(Chrysopyxis) or transverse division, and the swarming 
of one of the daughter-individuals (zoospore). Division 
may also take place in a motionless stage (pulmella-staye). 

Order 3. Dinobryinaceae. The individuals are ori- 
ginally attached, uni- or multi-cellular ; each individual 
cell is distinctly contractile, and fixed at the bottom of a 
cup-shaped, open envelope. Cilia 2, but of unequal 
length. Asexual reproduction by zoospores, which are 
formed by straight or oblique longitudinal division of 
the mother-cell, during a palmella-stage which is pro- 
duced in the winter aplauospores. Epipyxls, Dinolryon. 

FIG. 12. Chrsopyx-is 
m envelope, EC 
ckromatophure, cv 
contractile vacuole. 


Order 4. Hydruracese. The individuals are attached, without cilia, multi- 
cellular, branched, and with apical growth. The cells are spherical, but in 
the final stage almost spindle-shaped, and embedded in large masses of mucilage. 
Asexual reproduction by zoospores which are tetrahedric, with 1 cilia, and by 
resting akinetes. Hydrurus is most common in mountain brooks. 

Class -2. Dinoflagellata. 

The individuals are of a very variable form, but always uni- 
cellular, and floating about in free condition. The cell is dorsi- 
ventral, bilateral, asymmetric and generally surrounded by a colour- 
less membrane, which has no silica embedded in it, but is formed 
of a substance allied to cellulose. The membrane, which exter- 
nally is provided with pores and raised borders, easily breaks up into 
irregularly- shaped pieces. In the forms which have longitudinal and 
cross furrows, two cilia are fixed where these cross each other, and 
project through a cleft in the membrane ; one of these cilia projects 
freely and is directed longitudinally to the front or to the rear, the 
other one stretches crosswise and lies close to the cell, often in a 
furrow (cross furrow). The chromatophores are coloured brown 
or green and may either be two parallel (ExuvieUa), or several 
radially placed, discs, which sometimes may coalesce and become a 
star-shaped chromatophore. The colouring material 1 (pyrrophyl) 
consists, in addition to a modification of chlorophyl, also of phyco- 
pyrrin and peridinin ; this colour is sometimes more or less masked 
by the products of assimilation which, consist of yellow, red or 
colourless oil (?) and starch. Cell-nucleus one : in Polydinida several 
nuclei are found ; contractile vacuoles many, which partly open in 
the cilia-cleft (Fig. 13 gs). In some an eye-spot, coloured red by 
hasmatochrome, is found. Pyrenoids occur perhaps in ExuvieUa 
and AmpTiidinium. 

THE REPRODUCTION takes place as far as is known at present, 
only by division. This, in many salt water forms, may take 
place in the swarming condition, and, in that case, is always 
parallel to the longitudinal axis. The daughter-individuals, each 
of which retains half of the original shell, sometimes do not separ- 
ate at once from each other, and thus chains (e.g. in Ceratium} of 
several connected individuals may be formed. In others, the 
division occurs after the cilia have been thrown off and the cell-con- 
tents rounded. The daughter-cells then adopt entirely new cell-walls. 
A palmella-stage (motionless division-stage) sometimes appears to 



take place, and also aplanospores (?) with one or two horn-like 
elongations {e.g. in Peridinium cinctum and P. tabulatum) ; at 
germination one, or after division, two or more, new individuals 
may be formed. 

Sexual reproduction has not been observed with certainty. 

The Dinoflagellata move forward or backward, turning round 
their longitudinal axes ; in their motion they are influenced by 
the action of light. The motion possibly may be produced 
only by the transverse cilium, which vibrates rapidly ; whilst the 
longitudinal cilium moves slowly, and is supposed to serve mainly 
as a steering apparatus. They live principally in salt water, but 
also in fresh. 

Besides the coloured forms, which 
are able to make their own organic 
compounds by the splitting up of the 
carbonic acid contained in the water, 
there are a few colourless forms (e.g. 
Gymnodinium su'raZe),orsuch as do not 
possess chromatophores (Polykrikos) ; 
these appear to live saprophytically, 
and may be able to absorb solid bodies 
with which they come in contact. 

Dinoflagellata occur in the "Plank- 
ton " of the open sea, where they form 
together with Diatomacea? the basis 
for the animal life. It is known with 
certainty that some salt water forms 
(like the Noctiluca, which belongs -to 
the animal kingdom and to which 
they are perhaps related) produce 
light, known as phosphorescence. 

FIG. 13. A and B Gltnodiniuin 
cinctum. A Seen from the ventral 
side, B from behind ; fg transverse 
cilium ; g longitudinal cilium ; ch 
chromatophores ; a starch ; 11 cell- 
nucleus ; 13 vacuole ; oc eye-spot ; C 
Ceratium tetraceros from the ventral 
side ; i- the right, I the posterior 
horn ; If longitudinal furrow ; gs 
cilium-clef t ; D vacuole ; g longitudi- 
nal cilium. (A and B mag. 450 
times, C 337 times.) 

Dinqflagellata (Peridlnea, Cilioflagellata) are allied through their lowest form 
(Exuoiella) to the Syugeneticas and especially to the order Chrvsompnadinaceae. 
They may be divided into three orders. 

Order 1. Adinida. Without transverse or longitudinal furrows, but en- 
closed in two shells, and with two parallel chromatophores in each cell. 
Exuviella, Prorocentrum. 

Order 2. Dinifera. With tranverse and generally longitudinal furrow. Many 
radially-placed, disc-formed chromatophores. The most common genera are 
Ceratium (Fig. IB), Peridinium, Glenodinium (Fig. 13), Gymnodinium, Dlnophysis. 

OrderS. Polydinida. With several transverse furrows, no chromatophores, 
and several cell-nuclei. Only one genus Polykrikos. 

W. B. C 


The order PoJydinida deviates in a high degree from the other Dinorlagellata, 
not only by its many tranverse furrows, each with its own transverse cilium, and 
by the absence of chromatopbores, but also in havjng several cell-nuclei and a 
kind of stinging capsule, which otherwise does not occur within the whole 
class. It may therefore be questionable whether this order should really be 
placed in the vegetable kingdom. 

Class 3. Diatomeae. 

The individuals each known as a frustule assume very various 
forms and may be unicellular or multicellular, but present no 
differentiation ; many similar cells may be connected in chains, 
embedded in mucilaginous masses, or attached to mucilaginous 
stalks. The cells are bi-lateral or centric, often asymmetrical, 
slightly dorsiventral and have no cilia ; those living in the free 
condition have the power of sliding upon a firm substratum. The 
cell contains 1 cell-nucleus and 1-2 plate-shaped or several disc- 
shaped chromatophores. The colouring material " Melinophyl " 
contains, in addition to a modification of chlorophyl, a brown 
colouring matter, diatomin. 1 or 2 pyrenoids sometimes occur. 
Starch is wanting and the first product of assimilation appears to 
be a kind of oil (?). 

The cell-walls are impregnated with silica to such a degree that 
they are imperishable and are therefore able to contribute in a 
great measure to the formation of the earth's crust. The structure 
of their cell-wall is most peculiar and differs from all other plants 
(except certain Desmidiaceoe) ; it does not consist of a single piece 
but is made up of two the " shells " (compare Exuviella and 
Prorocentrum among the Dinoflagellata) which are fitted into each 
other, one being 1 a little larger than the other and embracing its 
edge, like a box with its lid (Fig. 14 B). The two parts which cor- 
respond to the bottom and lid of the box are known as valves. 
Along the central line of the valves a longitudinal rib may often 
be found, interrupted at its centre by a small cleft (perhaps 
homologous with the cilia-cleft of the Dinoflagellata), through 
which the protoplasm is enabled to communicate with the exterior 
(Fig. 14 A). It is principally by reason of the valves, which bear 
numerous fine, transverse ribs, striae or warts, etc. (Figs. 14, 15, 17), 
that the Diatomete have become so well known and employed as 
test objects in microscopical science. When the division takes 
place, the two shells are separated a little from each other, and 
after the cell-contents have divided into two masses, two new 
shells are formed, one fitting into the larger valve, the other one 




into the smaller valve of the original frustule. The latter cell 
(frustule) is thus, upon the whole, ^ -g 

smaller than the mother-cell, and as 
the cells do not increase in size, some 
frustules are smaller than the ones 
from which they are derived, and thus, 
by repeated divisions, it follows that 
smaller and smaller frustules are pro- 
duced. This continued diminution in 
size is, however, compensated for by 
the formation, when the cells have 
been reduced to a certain minimum, 
of auxospores, 2-3 times larger. These 
may either be formed asexually by the 
protoplasm of a cell increasing, round- 
ing off and surrounding itself with a 
new wall (e.y. Melosira} or after con- 
jugation, which may take place with 
various modifications: 1. Two indi- 
viduals unite after the secretion of a 
quantity of mucilage, and the valves 
then commence to separate from each 
other, on the side which the two indi- FlG - ^--P^nuiaria .- B, from 

. , . the edge, shows the valves fitting 

viduals turn towards each other. The together; .4, a valve. 

FIG. 15. Various Diatomacese. A Diatoma vulgare. B Tubellaria flocculosa. Navicula 
tumida (lateral views). D Gomphonema const ric twin, (lateral views). E Navicula u-estll 
(lateral views). 

protoplasmic bodies now release themselves from their cell-wall, 
and each rounds off to form an ellipsoidal mass ; these two pro- 


toplasmic masses (gametes) coalesce to form a zygote, tlie cell- 
nuclei and chromatophores also fusing together. The zygote in- 
creases in size, and surrounds itself with a firm, smooth, siliceous 
wall the perizonium. The auxospores, whichever way they arise, 
are not resting stages. The germination of the zygote com- 
mences by the protoplasm withdrawing itself slightly from the 
cell-wall and constructing first the larger valve, and later on 
the smaller one ; finally the membrane of the zygote bursts (e.g. 
Himantidiuni). 2. The conjugation occurs in a similar manner, 
but the protoplasm of the cells divides transversely before 
conjugation into two daughtei'-cells. Those lying opposite one 
another conjugate (Fig. 16) and form two zygotes. The for- 
mation of the perizonium, and germination take place as in the 
preceding instance (e.g. Epithemia). 3. Two cells place them- 
selves parallel to each other, and each of the two cell-contents, 


FIG. 16. Conjugation of Cijinbella variabilis. A, The protoplasm in the tvro cells has 
divided into two masses ; B these masses coalesce in pairs; the cells (B C) enclosed in a 
mucilaginous matrix. C D Auxospores and their formation. 

without coalescing, becomes an auxospore. The formation of 
the wall takes place as in the preceding case. This is found in 
the Naviculege, Cymbellere, the Gomphonemeae (e.g. FruStulia, 

The Diatomacese may be found in salt as well as in fresh water 
(often in such masses that the colour of the water or mud becomes 
yellow or brown ; in the same manner the genera Chcetoceros, 
Rhizosolenia, Coscinodiscus, and several others, form large slime- 
masses, " Plankton " on the surface of the sea), on damp soil and 
in dust blown by the wind. They occur as fossils in the recent 
formations, often in large deposits (siliceous earth, mountain 
meal), as in the cement lime in Jutland, the alluvial deposits 
beneath Berlin, in clay strata beneath peat bogs, in guano, etc. 


These accumulations of fossilized diatoms are used in the manu- 
facture of dynamite and in various manufactures. 

The Diatomaceae appear nearest to, and must be placed as a 
group co-ordinate with the Dinoflagellata, as they doubtless may 
be supposed to derive their origin from forms resembling Exuviella, 
and to have lost the cilia. The resemblances to the Desmidiaceo? 
which are striking in many respects, can only be conceived as 
analogies, and cannot be founded upon homologies, and it is 
therefore impossible to regard them as proof of genetic relation- 
ship. The family contains only one order. 

FIG. 17. Various Diatome. A Synedra radians. B EjntJiewu'a turgida (from the two 
different sides). C Cymbella cuspidata. D Cocconeis pediculus (on the right several situated 
on a portion of a plant, oil the left a single one more highly magnified). 

Order 1. Diatomacese. This order may be divided into two 
sub-orders, viz. 

Sub-Order 1. Placochromaticae. The chromatophores are 
discoid, large, 1 or 2 in each cell ; the structure of the valves 
is bilateral and always without reticulate markings. The follow- 
ing groups belong to this sub-order : Gomphonemex, Cynibellea>, 
Amphorece, Achnanthece, Cocconeidea 3 , Naviculece, Amphiplenrew, Phi- 
yiotropidea', Amphitropidece, Nitzchiece, Surirayece, and Eunotiece. 

Sub-Order 2. Coccochromaticse. The chromatophores are 
granular, small and many in each cell. The structure of the cells 
is zygomorphic or centric, often with reticulate markings. The 
following groups belong to this sub-order : Fragilariece, Meridiece, 
Tabellariece, Licmophorece, Biddulphiece, AngulifercB, Eupodiscece, 
Coscinodiscece and Melosirece. 


Class 4. Schizophyta, Fission-Algae. 

The individuals are 1 many celled ; the thallus consists in 
many of a single cell, in others of chains of cells, the cells dividing 
in only one definite direction (Figs. 18, 21). In certain Fission-Alga? 
the cell-chain branches (Fig. 30) and a difference between the an- 
terior and the posterior ends of the chain is marked ; in some, the 
cells may be united into the form of flat plates by the cell-division 
taking place in two directions ; and in others into somewhat cubi- 
cal masses, or rounded lumps of a less decided form, by the divisions 
taking place in three directions ; or less defined masses may be 
formed by the divisions taking place in all possible directions. 

The cell- walls rarely contain cellulose, they often swell con- 
siderably (Figs. 20, 22), and show distinct stratifications, or they 
are almost completely changed into a mucilaginous mass in which 
the protoplasts are embedded, e.g. in Nostoc (Fig. 22), and in the 
"Zoogloea" stage of the Bacteria (Fig. 27). Sexual reproduction 
is wanting. Vegetative reproduction by division and the separa- 
tion of the divisional products by the splitting of the cell- wall or 
its becoming mucilaginous ; among the Nostocacea?, Lyngbyaceie, 
Scytonemacese, etc., " Hormogonia " are found ; in Ghamcesiphon 
and others single reproductive akinetesare formed. Many Fission- 
Alga? conclude the growing period by the formation of resting 
akinetes or aplanospores. 

The Schizophyta may be divided into 2 families : 



Family 1. Schizophyceae,* Blue-Green Algse. 
All the Blue-green Alga? are able to assimilate carbon by means 
of a colouring material containing chlorophyll (cyanophyll) ; but 
the chlorophyll in this substance is masked by a blue (phycocyan), 
or red (phycoerythrin, e.g. in Trichodesmium erythrceum in the Red 
Sea) colouring matter which may be extracted from them in cold 
water after death. The colouring matter, in most of them, per- 
meates the whole of the protoplasm (excepting the cell-nucleus), 
but in a few (e.g. Glaucocys/is, Phragmonema), slightly developed 
chromatophores are to be found. Where the cells are united into 
filaments (cell-chains) a differentiation into apex and base (Rivu- 
lariacece) may take place, and also between ordinary vegetative 
cells and heterocysts; these latter cannot divide, and are dis- 

* Myxophyceas, Cyauophycere. 


tinguished from the ordinary vegetative cells (Fig. 22 h~) by their 
larger size, yellow colour, and poverty of contents. Branching 
sometimes occurs and is either true or spurious. 

FIG. IS. Microcoleus lyngbyanus; a portion of a filament, the thick sheath encloses only 
one cell-chain ; in one place a cell is drawn out by the movement of the cell-chain; I the 
cell-chain has divided into two parts (" hormongonia ") which commence to separate 
from each other. 

The cell-chain in the spurious branching divides into two parts, 
of which either one or both grow beyond the place of division 
(Fig. 18) and often out to both sides (e.g. Scytonema), the divisions 
however, always take place transversely to the longitudinal 
direction of the cell-chain. In the true branching a cell elon- 
gates in the direction transverse to the cell-chain, and the division 
then takes place nearly at right angles to the former direction 
( Sirosiphoniacece). 

FIG. 19. Cj/Iindi'ugf)<!rmu(ii majws : a resting akinete with heterocyst : b-d germinating 
stages of a resting akinete ; e filament with two heterocysts and the formation of new 
akinetes; / part of a filament with a heterocysfc, and mature resting akinete. 

Cilia are wanting, but the filaments are sometimes self-motile 
(e.g. hormogonia in Nostoc) arid many partly turn round their axes, 
partly slide forward or backward (Oscillaria). 

Reproduction takes place by spores and horraogonia in addition 



to simple cell-division. Hormogonia are peculiar fragments of a 
cell-chain capable of motion, and often exhibit a vigorous motion 
in the sheath, until at last they escape and grow into a new- 
individual (Fig. 18). The spores are reproductive akinetes 
(Chamcesiphon, etc.) or resting akinetes ; these latter arise by the 
vegetative cells enlarging and constructing a thick cell-wall (Fig. 
19 ef). On germination, this cell-wall bursts and the new cell-chain 
elongates in the same longitudinal direction as before (Fig. 19 6 c). 
Many ( OsciHaria) may however winter in their ordinary vege- 
tative stage. Aplanospores are wanting. 

The Fission- Algae are very prevalent in fresh water and on 
damp soil, less so in salt water ; they also often occur in water 
which abounds in decaying matter. Some are found in warm 
springs with a temperature as high as 50 C. 

The Family may be divided into 2 sub-families : 

1. HOMOCYSTE.S; (heterocysts are wanting) : 
Lyngbyacece and Chamcesiphonacece. 

2. HETEROCYSTEJI (heterocysts present) : Nostocacece, 
ariacew, Scytonemacece and Sirosiphoniacece. 

Order 1. Chroococcaceae. The individuals are 1 many- 
celled, but all the cells are uniform, 




united to form plates or irregular 

masses, often surrounded by a mu- 
cilaginous cell-wall, but never forming 
cell-chains. Multiplication by division 
and sometimes by resting akinetes, but 
reproductive akinetes are wanting. 
Chroococcus, ApJianocapsa, Gloeocapsa 
( Fig. 20) , Ccelosphaerium, Merism opedium , 
Glaucocystis, Oitcobyrsa, Poli/cystis, Gom- 

Order 2. Lyngbyaceae (Oscil- 
The cells are discoid (Fig. 21), united to straight or 

spirally twisted, free filaments, 
which are unbranched, or with 
spurious branching. The ends 
of the cell-chains are .similar. 
Heterocysts absent. Reproduc- 
tion by synakinetes, resting 
akinetes are wanting. Oscillaria (Fig. 21), Spindina, Lyvgbya, 
Microcoleus, Symploca, Plectonema. 

Fie. 20. Glceocapsa atrata: A, 
S, C, D, E various stages of 


FIG. 21. Osi'iVfori'n; n terminal, l> central 
poitinii of a filament. 


Order 3. Chamaesiphonaceae. The individuals are 1- 
many-celled, attached, unbranched filaments with differentiation 
into apex and base, without heterocysts. Multiplication by re- 
productive akinetes; resting akinetes are wanting. Dermocarpa, 
Clastidium, Chamcesiphon, Godlewskia, Phragmonema. 

Order 4. Nostocaceae. The individuals are formed of mul- 
ticellular, unbranched filaments, without differentiation into apex 
and base ; heterocysts present. Reproduction by synakinetes and 
resting akinetes. 

Some genera are not mucilaginous, e.g. Cylindrospermum (Fig. 
19). The cell-chains in others, e.g. Nostoc, wind in between one 

FIG. 22. Nostoc verrucostim. A The plant in its natural size ; an irregularly folded 
jelly-like mass. B One of the cell-chains enlarged, with its heterocysts (ft), embedded in 
its mucilaginous sheath. 

another and are embedded in large structureless jelly-like masses, 
which may attain the size of a plum or even larger (Fig. 22) ; 
sometimes they are found floating in the water, sometimes 
attached to other bodies. Other genera as follows : Aphanizo- 
menon and Anabc&na (in lakes and smaller pieces of water) ; 
Nodularia is partly pelagic. Some occur in the intercellular 
spaces of higher plants, thus Nostuc-forms are found in Anthocerus, 
Blasia, Sphagnum, Lemna, and in the roots of Cijcas and Gunnera ; 
Anabvena in Azolla. 

Order 5. Rivulariaceae. The individuals are multicellnlar 
filaments, with differentiation into apex and base ; spurious 
branching, and a heterocyst at the base of each filament. Re- 
production by synakinetes and resting akinetes, rarely by simple 
reproductive akinetes. Rivulama, Glceotrichia, Isactis, Caluthrix. 

Order 6. Scytonemaceae. The individuals are formed of 
mnlticellular filaments with no longitudinal division ; differen- 
tiation into apex and base very slight or altogether absent ; 



branching spurious ; heterocysts present. Reproduction by syna- 
kinetes, rarely by resting akinetes and ordinary reproductive 
akinetes. Tolypothrix, Scytonema, Hassalia, Microchcete. 

Order 7. Sirosiphoniaceae. The individuals are formed of 
multicellular threads with longitudinal divisions ; true branching 
and heterocysts, and often distinct differentiation into apex and 
base. Reproduction by synakinetes, rarely by resting akinetes 
and ordinary reproductive akinetes. Hapalosiphon, Stigonema, 
Capsosira, Kostocopsis, Mas/igocolens. 

Family 2. Bacteria.* 

The Bacteria (also known as Schizomycetes, and Fission-Fungi) 
are the smallest known organisms, and form a parallel group to 
the Blue-green Algae, but separated from these Algae by the 
absence of their colouring material ; chlorophyll is only found in 
a few Bacteria. 

The various forms under which the vegetative condition of the 
Bacteria appear, are termed as follows : 

1. GLOBULAR FORMS, cocci (Figs. 27, 30 c) : spherical or ellip- 
soidal, single cells, which, howevei-, are usually loosely massed 
together and generally termed "Micrococci." 

2. ROD-LIKE FORMS : more or less elongated bodies ; the shorter 
forms have been styled "Bacterium" (in the narrower sense of the 
word), and the term "Bacillus " has been applied to longer forms 

which are straight 
and cylindrical (Figs. 
28, 29, 30 E). 

FORMS : unbranched, 
long, round filaments, 
resembling 1 those of 
Oscillaria, are pos- 
sessed by Leptothrix 
(very thin, non-granu- 
lar filaments ; Fig. 30 
^4, the small filaments) 
and Beggiatoa (thicker 
filaments, with strong, refractile grains or drops of sulphur (Fig. 

FIG. 23. Spirillum sanguineutn. Four specimens. One 
has two cilia at the same end, the sulphur grains are seen 

* The Bacteria are more usually included under Fungi. It seems better, how- 
ever, to place them under the Algae in a separate class with the Suhizophyceae. 



31); often self-motile). Branched filaments, with false brandling 
like many Scytonemacece, are found in Cladothrix (Fig. 30 B, G}. 

4. SPIRAL FORMS : Rod-like or filamentous bodies, which more 
or less strongly resemble a corkscrew with a spiral rising to the 
left. In general these are termed Spirilla (Fig 23) ; very attenu- 
ated spirals, Vibriones (standing next to Fig. 30 M) ; if the filaments 
are slender and flexible with a closely wound spiral, 

(Fig. 24). 

FORM, consisting of rounded 
cells arranged in one plane, 
generally in groups of four, and 
produced by divisions perpen- 
dicular to each other. 

6. The SARCINA-FORM, con- 
sisting of roundish cells which 
are produced by cellular divi- 
sion in all the three directions 
of space, united into globular 
or ovoid masses ("parcels") 
e.g. Sarcina ventriculi (Figs. 
25, 26). 

All Bacteria are unicellular. In the case of the micrococci this 
is self-evident, but in the " rod," " thread," and " spiral " Bacteria, 
very often numerous cells remain united together and their indi- 
vidual elements can only be recognised by the use of special re- 

FIG. 24. Spirochxte obermeieri, in active 
motion (b)aiid shortly before the termination 
of the fever (c); o blood corpuscles. 



FIG. 25. Soi-ciua ventnculi. One sur- 
face only is generally seen. Those cells 
which are drawn with double contour are 
seen with the correct focus, and more 
distinctly than those cells lying deeper 
drawn with single contour. 

FIG. 26. Sarchia mmuta : a-d succes- 
sive stages of one individual (from 4-10 
p.m.) ; / an individual of 32 cells. 

The condition termed " Zooglcea," which reminds us of Nostoc, is 
produced by the cells becoming strongly mucilaginous. A number 
of individuals in active division are found embedded in a mass of 
mucilage, which either contains, only one, or sometimes more, of 


the above-named forms. The individuals may eventually swarm 
out and continue their development in an isolated condition. 
Such mucilaginous masses occur especially upon moist vegetables 
(potatoes, etc.), on the surface of fluids with decaying raw or 
cooked materials, etc. The mucilaginous envelope is thrown into 
folds when the Bacteria, with their mucilaginous cell-walls, 
multiply so rapidly that there is no more room on the surface of 
the fluid. 

The cells of the Bacteria are constructed like other plant-cells 
in so far as their diminutive size has allowed us to observe them. 
The cell-wall only exceptionally shows the reactions of cellulose 
(in Sarcina, Leuconostoc ; also in a Vinegar-bacterium, Bacterium 
xylinum) ; a mucilaginous external layer is always present. The 
body of the cell mostly appears to be an uniform or finely granu- 
lated protoplasm. Very few species (e.g. Bacillus virens) contain 
chlorophyll ; others are coloured red (purple sulphur Bacteria) ; 
the majority are colourless. Bacillus amyloLacter shows a reaction 
of a starch-like material when treated with iodine before the 
spore-formation. Some Bacteria contain sulphur (see p. 37). 
The body, which has been described as a cell-nucleus, is still of a 
doubtful nature. 

Artificial colourings with aniline dyes (especially methyl-violet, 
gentian-violet, methylene-blue, fuchsin, Bismarck-brown and 
Vesuvin) play an important part in the investigations of 

MOVEMENT. Many Bacteria are self-motile; the long filaments of 
Beggiatoa exhibit movements resembling those of Oscillaria. In 
many motile forms the presence of cilia orflagella has been proved 
by the use of stains ; many forms have one, others several cilia 
attached at one or both ends (Fig. 23) or distributed ii-regularly 
over the whole body ; the cilia are apparently elongations of the 
mucilaginous covering and not, as in the other Algas of the proto- 
plasm. In Spirochcete the movement is produced by the flexibility 
of the cell itself. Generally speaking, the motion resembles that 
of swarm-cells (i.e. rotation round the long axis and movement in 
irregular paths) ; but either end has an equal power of proceeding 

The swarming motion must not be confounded with the hopping motion of the 
very minute particles under the microscope (Browiiian movement). 

VEGETATIVE REPRODUCTION takes place by continued transverse 



division ; hence the name " Fission-Fungi " or " Fission-Algae," 
has been applied to the Bacteria. 

SPORES. The spores are probably developed in two ways. In 
the ENDOSPOROUS species (Figs. 28, 29), the spore arises as a new 
cell inside the mother-cell. The spores are strongly refractile, 
smaller than the mother-cell, and may be compared to the 
aplanospores of other Algne. In addition to these there are the 
ARTHROSPOROUS species in which the cells, just as in Nostoc and 
other Blue-green Algge, assume the properties of spores without 
previously undergoing an endogenous new construction, and are 
able to germinate and form new vegetative generations (Fig. 27). 
The formation of spores very often commences when the vegetative 
development begins to be restricted. 

FIG. 27. Leuconostoc mcsentevioides : a a zoogloea, natural size ; b cross section of 
zoogloea ; c filaments with spores ; d mature spores ; e-i successive stages of germina- 
tion ; in e portions of the ruptured spore-wall are seen on the external side of the muci- 
laginous covering, (b-i magnified 520.) 

The spores germinate as in Nostoc by the bursting of the external 
layer of the cell-wall, either by a transverse or longitudinal cleft, 
but always in the same way, in the same species (Fig. 28, example 
of transverse cleft). 

DISTRIBUTION. Bacteria and their germs capable of development, 
are found everywhere, in the air (dust), in surface water, and 
in the superficial layers of the soil. The number varies very 
much in accordance with the nature of the place, season, etc. 
They enter, together with air and food, into healthy animals and 
occur always in their alimentary tract. 



GROWTH AND REPRODUCTION depend upon the conditions of 
temperature. There is a certain minimum, optimum and maxi- 
mum for each species ; for instance (in degrees Centigrade) 

VIG. 23. Bacillus megaterium : a outline of a 
living, vegetative cell-rod ; b a living, motile, pair of 
rods ; p a similar 4-celled rod after the effects of 
iodine alcohol ; c a 5-celled rod in the first stages of 
spore-formation ; d-f successive stages of spore- 
formation in one and the same pair of rods (in the 
course of an afternoon) ; r a rod with mature spores ; 
g'-;/ 3 three stages of a 5-celled rod, with spores sown 
in nutritive solution ; Ji'-fi 3 , i,fe, I stages of germina- 
tion ; m a rod in the act of transverse division, 
grown out from a spore which had been sown eight 
hours previously. (After de Bary ; a mag. 250, the 
other figures 600 times). 

Bacillus subtilis 

B. anlhracis 

Spirillum cholerce asiaticce 

Bacterium tuberculosis 

+ 6 



c. 30 

28 37-38 

FIG. 29. Bacillus amj/lobacter. 
Motile rods, partly cylindrical 
and without spores, partly 
swollen into various special 
shapes and wilh spore-forma- 
tion in the swelling, s Mature 
spore, with thick mucilaginous 
envelope. (After de Bary ; 
mag. 600 times, with the excep- 
tion of s, which is more highly 

+ 50 


40 (but grows only 

feebly if under 


The functions of life cease on a slight excess of the maximum 
or minimum temperature, numbness setting in when either of 
these limits is passed. Crenothrix-ila.resLda provided with muci- 
laginous envelopes may, according to Zopf, sustain a tempera- 
ture of 10. Some Bacteria are said to be able to resist the 
exposure to as low a temperature as 110 for a short time. It is 
not known at what degree of cold the death of the Bacteria oc- 
curs : the greatest degree of heat which the vegetative cells can 


withstand is about the same as that for other vegetative plant- 
cells, namely, about 50-60 C. Certain Bacteria, e.g. B. tliermo- 
philus, grow and thrive vigorously at 70 C. Many spores, on the 
contrary, are able to bear far higher temperatures (in several 
species a temperature for some duration of above 100, those of 
Bacillus subtilis, for instance, can withstand for hours a tempera- 
ture of 100 in nutrient solutions ; the spores remain capable of 
development after exposure to a dry heat of 123 C.). 

The Desiccation of the air, if prolonged, kills many forms when 
in the vegetative condition. The spores however can bear a much 
longer period of dryness, some even several years. 

OXYGEN. Some species cannot live without a supply of free 
oxygen (Aerobic), e.g. the Vinegar-bacteria, the Hay-bacilli, the 
Anthrax-bacilli, the Cholera- Microspira. Other species again thrive 
vigorously without supply of free oxygen, and are even checked in 
their development by the admission of air (Anaerobic), e.g. the 
butyric acid Bacterium (Glostridium butyricum = Bacillus amy- 
lobacter). A distinction may be drawn between obligate and 
facultative aerobics and obligate and facultative anaerobics. 
Several Bacteria, producing fermentation, may grow without the 
aid of oxygen when they are living in a solution in which they 
can produce fermentation ; but, if this is not the case, they can 
only grow when a supply of oxygen is available. A great 
number of the pathogenic Bacteria belong to the facultative 

A luminous Bacterium (Bacillus phosphorescens) which in the 
presence of a supply of oxygen gives a bluish-white light, has 
been found in sea- water. Phosphorescent Bacteria have fre- 
quently been observed upon decaying sea-fish, as well as on the 
flesh of other animals ; by transferring the Bacteria from cod fish 
to beef, etc., the latter may be made luminous. 

Organic carbon compounds are indispensable for all Bacteria, 
(except, as it appears, for the nitrifying organisms), as they can 
only obtain the necessary supplies of carbon from this source. The 
supplies of nitrogen, which also they cannot do without, can be ob- 
tained equally as well from organic compounds as from inorganic 
salts, such as saltpetre or ammonia-compounds. The various "ash- 
constituents " are also essential for their nourishment. 

While Moulds and Yeast-Fungi grow best in an acid substratum, 
the Bacteria, on the other hand, generally thrive best in a neutral 
or slightly alkaline one. 


In sterilization, disinfection, and antisepsis, means are employed 
by which the Bacteria are killed, or. checked in their development, 
for instance, by heat (ignition, cooking, hot vapours, hot air, etc.), 
or poisons (acids, corrosive sublimate). The process of preserv- 
ing articles of food, in which they are boiled and then hermeti- 
cally sealed, aims at destroying the Bacteria, or the spores of 
those which already may be present in them, and excluding 
all others. 

As the Bacteria are unable to assimilate carbon from the car- 
bonic acid of the air, but must obtain it from the carbon-com- 
pounds already in existence in the organic world, they are either 
saprophytes or parasites. Some are exclusively either the one or 
the other, obligate saprophytes or parasites. But there are 
transitional forms among them, some of which are at ordinary 
times saprophytes, but may, when occasion offers, complete their 
development wholly or partly as parasites facultative parasites ; 
others are generally parasitic, but may also pass certain stages of 
development as saprophytes facultative saprophytes. 

All chlorophyll-free organisms act in a transforming and dis- 
turbing manner on the organic compounds from which they obtain 
their nourishment, and while they themselves grow and multiply, 
they produce, each after its kind, compounds of a less degree of 
complexity, i.e. they produce fermentation, putrefaction, sometimes 
the formation of poisons, and in living beings often disease. 

Those organisms which produce fermentation are called ferments ; 
this word, however, is also employed for similar transformations 
in purely chemical materials (inorganic ferments or enzymes). 
Many organic (" living ") ferments, among which are Yeast- 
cells and Bactei'ia, give off during their development certain 
inorganic and soluble ferments (enzymes) which may produce 
other transformations without themselves being changed. Dif- 
ferent organisms may produce in the same substratum different 
kinds of transformation ; alcoholic fermentation may for instance 
be produced by different species of Fungi, but in different pro- 
portions, and the same species produces in different substrata, 
different transformations (e.g. the Vinegar-bacteria oxydize diluted 
alcohol to vinegar, and eventually to carbonic acid and water). 

In the study of Bacteria it is absolutely necessary to sterilize the vessels 
employed in cultivation, the apparatus, and nutrient solutions, i.e. to free them 
from Bacteria germs and also to preserve the cultures from the intrusion of any 
foreign germs ("pure-cultures"). A firm, transparent, nutritive medium is 


frequently employed. This may be prepared by adding to tbe nutrient solu- 
tions (broth) either gelatine, or when the Bacteria are to be cultivated at 
blood-heat serum of sheep's or calf's blood, agar-agar or carragen ; serum 
alone may in itself serve as a nutrient medium. The so-called " plate-cul- 
tures " are frequently employed, i.e. the germs are isolated by shaking them 
with the melted liquid nutrient gelatine, which is then spread on a glass plate 
and allowed to coagulate ; when later on the individual germs grow into 
colonies, these remain separate in the solid substratum and it is easy to pursue 
their further development. Similar plate-cultures may also be cultivated in 
test-tubes and on microscopic slides. The slides and glass plates must be 
placed in " moist chambers" free from Bacteria. By sowing a few cells (if pos- 
sible one) by means of a fine platinum wire, pure cultures for further investi- 
gation may be obtained. 

In order to prove the relationship between pathogenic Bacteria and certain 
diseases, the experimental production of pathogenic Bacteria by the inoculation 
of Bacteria from pure cultures into healthy animals, is very important. 

It has not so far been possible to establish a classification of the 
Bacteria, as the life-history of many species has not yet been 
sufficiently investigated. 1 The opinions of botanists are at 
variance, in many cases, with regard to the forms of growth of a 
particular kind. Some species are pleomorphic (many- formed) 
while others possess only one form. 

The following Bacteria are Saprophytes : 

Gladothrix dichotonia is common in stagnant and running water 
which is impregnated with or-ganic matter; the cell-chains have 
false branching. According to Zopf, Leptothrix ochracea is one of 
the forms of this species which, in water containing ferrous iron 
(e.g. as FeC0 3 ), regularly embeds ferric-oxide in its sheath by means 
of the activity of the protoplasm. Leptothrix ochracea and other 
Iron-bacteria, according to Winogradsky (1888), do not continue 
their growth in water free from protoxide of iron ; while they 
multiply enormously in water which contains this salt of iron. 
The large masses of ochre-coloured slime, found in meadows, bogs, 
and lakes, are probably due to the activity of the Iron-bacteria. 

Those forms which, according to Zopf's views, represent the 
forms of development of Cladothrix dichotoma are placed together 
in Fig. 30. A represents a group of plants, seventy times 
magnified, attached to a Vaucheria. The largest one is branched 
like a tree, with branches of ordinary form ; a specimen with 
spirally twisted branches is seen to the right of the figure, at 
the lower part some small LeptotJirix-like forms. B shows the 

1 See Marshall Ward, " On the Characters or Marks employed for Classifying 
the Schizomyceles," Annals of Botany, 1892. 

W. B. D 



manner of branching and an incipient Cocctts-formation. C a 
Coccus-mass whose exit from the sheath has been observed. D the 

FIG. 3/0.Cladothrix dicliotoma. 

same mass as G after the course of a day, the Cocci having 
turned into rods. E a group of Cocci in which some have deve- 
loped into shorter or longer rods. F one of these rods before 


and after treatment witli picric acid, which causes the chain-like 
structure to become apparent. G a portion of a plant with con- 
spicuous sheath, two lateral branches are being- formed. JET part 
of a plant, whose cells have divided and form Cocci. The original 
form of the cells in which the Cocci are embedded may still be 
recognised. I. Leptothrix-Gla,m.Qa.ta with conspicuous mucilaginous 
sheath, from which a series of rods is about to emerge ; the rod 
near the bottom is dead, and has remained lying in the sheath. 
K part of a plant which is forming Cocci, those at the top are in 
the zoogloea-stage, at the base they are elongating to form rods 
and Leptothrix-fil&meui,B. L a portion of a branched Cladothrix, 
which divides into motile Bacillus-forms ; the rays at the free 
ends indicate the currents which the cilia produce in the water. 
M a spirally-twisted, swarming filament, before and after division 
into halves. N part of a tree-like zoogloea with Cocci and short 
rods. All of these spirilla, zoogloea, etc., which Zopf has con- 
nected with Clad, dichotoma, are according to Winogradsky, inde- 
pendent organisms. 

Micrococcus urece produces urinal fermentation (transformation of 
urinal matter into ammonium carbonate) ; aerobic ; round cells 
generally united to form bent chains or a zoogloea. Several other 
kinds of Bacteria have the same action as this one : in damp 
soil containing ammonia-compounds, saltpetre-formations are pro- 
duced by M. nitrificans and several different kinds of Bacteria. 

Micrococcus prodigiosus is found on articles of food containing 
starch ; " bleeding bread " is caused by this Bacterium, which has 
the power of forming a red pigment ; it also occurs in milk, and 
produces lactic acid. 

Leuconostoc mesenterioides is the frog-spawn Bacterium (Fig. 27) 
which is found in sugar manufactories, and has the power of 
producing a viscous fermentation in saccharine solutions which 
have been derived from plants, e.g. in beetroot-sugar manu- 
factories, where large accumulations of mucilage are formed at 
the expense of the sugar, with an evolution of carbonic acid. The 
cell-rows, resembling somewhat a pearl necklace, have thick 
mucilaginous cell-walls, and form white " Nostoc "-lumps. The 
mucilage eventually deliquesces and the cells separate from each 
other; arthrospores ? -Similar viscous deteriorations occur in 
beer and wine, which may then be drawn out into long, string 
like filaments " ropiness." 

Bacterium aceti, the Vinegar-bacterium, oxidizes alcohol into 


acetic acid (acetous-fermentation) and forms a greyish covering 
of Bacteria ("Vinegar-mother") on the surface of the liquid ; the 
acetic acid formed, becomes by continued oxidization by B. aceti, 
again transformed into carbonic acid and water. Aerobic; 
short cylindrical cells, often united into chains, or to form a 
zoogloaa ; sometimes also rod- and spindle-shaped. The Vinegar- 
bacteria and other kinds with ball- or rod-forms sometimes become 
swollen, spindle-shaped, or oval links ; they are supposed to be 
diseased forms 1 ("Involution-forms"). 

Bacillus laciictts (Bacterium acidi lactici, Zopf) is always found 
in milk which has stood for some time, and in sour foods (cabbage, 
cucumbers, etc.) ; it turns the milk sour by producing lactic acid 
fermentation in the sugar contained in the milk ; the lactic acid 
formed, eventually causes the coagulation of the casein. It re- 
sembles the Vinegar-bacteria, occurring as small cylindrical cells, 
rarely in short rows; not self-motile. Several other Bacteria appear 
to act in the same way, some occurring in the mouth of human 
beings ; some of these Bacteria give to butter its taste and flavour. 

The kefir-grains which are added to milk for the preparation of 
kefir, contain in large numbers a Bacterium (Dispora caucasica') 
in the zoogloea-form, a Yeast-fungus, and Bacilhis lacticus. Kefir 
is a somewhat alcoholic sour milk, rich in carbonic acid ; it is a 
beverage manufactured by the inhabitants of the Caucasus, from 
the milk of cows, goats, or sheep, and is sometimes used as a 
medicine. In the production of kefir, lactic acid fermentation 
takes place in one part of the sugar contained in the milk, and 
alcoholic fermentation in another part, and the casein which had 
become curdled is partially liquefied (peptonised) by an enzyme of 
a Zoogloea-bacterium. 

IlacUlns amylobacter (Bacillus buiyricus) , the Butyric-acid-bac- 
terium (Fig. 29), is a very common anaerobic which produces 
fermentation in sugar and lactic-acid salts, and whose principal 
product is butyric acid. It destroys articles of food and (together 
with other species) plays a part in the butyric acid fermentation 
Avhich is necessary in the making of cheese ; it is verv active 
wherever portions of plants are decaying, in destroying the cellu- 
lose in the cell-walls of herbaceous plants, and is thus useful in 
the preparation of flax and hemp. The cells are self-motile, 
generally cylindrical, sometimes united into short rows ; endo- 

1 According to Hausen these are net disease formr, bat occur regularly under 
certain conditions, e.g. temperature. 







sporous; the spore-forming cells swell, assume very different forms, 
and show granulose reaction. The germ-tube grows out in the 
direction of the long axis of the spore. 

Bacillus snbtilis, the Hay-bacillus, is developed in all decoctions 
of hay ; a slender, aerobic, self-motile Bacillus ; endosporous (aplano- 
spores) ; the spore-wall ruptures transversely on germination. 

Crenothrix kilhuiana occurs in the springs of many baths, in 
wells, in water or drain-pipes. 

Beggiatoa (parallel with the Blue-green Alga Oscillaria). Long 
filaments formed of cylindrical cells which are attached by one of 
the ends, but which are nearly always free when observed. The 
filaments, like those of Oscillaria, describe conical figures in their 
revolutions, the free filaments slide upwards and parallel with one 
another; sheaths are wanting ; strongly 

refractive sulphur drops are found in the *| ^ r\ 

interior. The Beggiatoas are the most W H y\ 

iriai IE a 

prevalent Sulphur-bacteria. They occur, 
very commonly in large numbers, wher- 
ever plant or animal remains are de- 
caying in water in which sulphuretted 
hydrogen is being formed ; thus, for ex- 
ample, B. alba (Fig. 31) occurs frequently 
as a white covering or slimy film on mud 
containing organic remains. B. mirabilis is 
remarkable for its size and its strong peristaltic 
movements. The Sulphur-bacteria oxidize 
the sulphuretted hydi-ogen, and accu- 
mulate sulphur in the shape of small 
granules of soft amorphic sulphur, which 
in the living cell never passes over into 
the crystalline state. They next oxidize 
this sulphur into sulphuric acid, which 
is immediately rendered neutral by ab- 
sorbed salts of calcium, and is given 
off in the form of a sulphate, thus CaC0 3 is principally changed 
into Ca SO 4 . In the absence of sulphur the nutritive processes 
are suspended, and consequently death occurs either sooner or 
later. The Sulphur- bacteria may exist and multiply in a fluid 
which only contains traces of organic matter, in which organisms 
devoid of chlorophyll are not able to exist. The Beggiatoas 
very frequently form white, bulky masses in sulphur wells and 



FIG. 31. Beggiatoa alba: a 
from a fluid containing abund- 
ance of sulphuretted hydrogen ; 
I) after lying 24 hours in a solu- 
tion devoid of sulphuretted 
hydrogen ; c after lying an 
additional -48 hours in a solution 
devoid of sulphuretted hydro- 
gen, by this means the trans- 
verse walls and vacuoles have 
become visible. 


in salt water, the traces of organic material which the sulphur 
water contains proving sufficient for them. The cellulose-fermenta- 
tion, to which the sulphur wells iu all probability owe their origin, mainly 
procures them suitable conditions for existence. The Ca C0 3 and H 2 S, 
formed during the cellulose fermentation by the reduction of CaS0 4 is 
again changed into CaSOj and CO 2 by the Sulphur-bacteria (Winogradsky, 
1887). Other Sulphur-bacteria, the so-called purple Sulphur-bacteria, e.ij. 
B. roseo-persicina, Spirillum sanguineum (Fig. 23), Bacterium sulfuratum, etc., 
have their protoplasm mixed with a red colouring matter (bacterio-purpurin) 
which, like chlorophyll, has the power, in the presence of light, of giving off 
oxygen (as proved by T. W. Englemann, 1888, in oxygen-sensitive Bacteria). 
The three purple Sulphur-bacteria mentioned, are, according to Winogradsky, 
not pleomorphic kinds but embrace numerous species. 

Many Spirilli (Spirillum tenue, S. nndula, S.plicatile, and others) 
are found prevalent in decaying liquids. 

Bacteria (especially Bacilli) are the cause of many substances 
emitting a foul odour, and of various changes in milk. 

Parasitic Bacteria live in other living organisms ; but the 
relation between "host" and parasite may vary in considerable 
degree. Some parasites do no injury to their host, others produce 
dangerous contagious diseases ; some choose only a special kind 
as host, others again live equally well in many different ones. 
There are further specific and individual differences with regard 
to the predisposition of the host, and every individual has not the 
same receptivity at all times. 

above mentioned saprophytes may also occur in the alimentary 
canal of human beings ; e.g., the Hay-bacillus, the Butyric-acid- 
bacillus, etc. ; but the gastric juice prevents the development of 
others, at all events in their vegetative condition. Sarcina ven- 
triatli, "packet-bacterium," is only known to occur in the stomach 
and intestines of human beings, and makes its appearance in cer- 
tain diseases of the stomach (dilation of the stomach, etc.) in great 
numbers, without, however, being the cause of the disease. It 
occurs in somewhat cubical masses of roundish cells (Fig. 25). 

LESS DANGEROUS PARASITES. In the mouth, especially between 
and on the teeth, a great many Bacteria are to be found (more 
than fifty species are known), e.g. Leptothrix buccalis (long, brittle, 
very thin filaments which are united into bundles), Micrococci in 
large lumps, Spirochtvte cohnii, etc. Some of them are known to 
be injurious, as they contribute in various ways to the decay of the 
teeth (caries dentiuvn) ; a Micrococcns, for instance, forms lactic acid 


in materials containing sugar and starch, and the acid dissolves 
the lime salts in the external layers of the teeth : those parts of 
the teeth thus deprived of lime are attacked by other Bacteria, and 
become dissolved. Inflammation in the tissues at the root of a 
tooth, is probably produced by septic materials which have been 
formed by Bacteria in the root-canal. 

DANGEROUS PARASITES. In a large number of the infections dis- 
eases of human beings and animals, it has been possible to prove 
that parasitic Bacteria have been the cause of the disease. Various 
pathogenic Bacteria of this nature, belonging to the coccus, rod, 
and spiral Bacteria groups, are mentioned in the following: 

Pathogenic Micrococci. Staphylococcus pyogenes aureus pro- 
duces abscesses of various natures (boils, suppurative processes in 
internal organs). The same effects are produced by 

Streptococcus pyogenes, which is the most frequent cause of malig- 
nant puerperal fever ; it is perhaps identical with 

Streptococcus erysipelatis, which is the cause of erysipelas in 
human beings. 

Diplococcus pneumonia (A. Friinkel) is the cause of pneumonia, 
and of the epidemic cerebro-spinal meningitis. 

Gonococcus (Neisser) is the cause of gonorrhea and inflammation 
of the eyes. 

Pathogenic Rod- Bacteria. Bacterium cholerce gallinarum, 
an aerobic, facultative parasite which produces fowl-cholera among 
poultry ; it is easily cultivated on various substrata as a sapro- 
phyte. The disease may be conveyed both through wounds and 
by food, and may also be communicated to mammals. 

Bacillus anthraci*, the Anthrax bacillus (Fig. 32), chiefly attacks 
mammals, especially herbivorous animals (house mice, guinea-pigs, 
rabbits, sheep, cattle), in a less degree omnivorous animals (includ- 
ing human beings), and in a still less degree the Carnivores. 
Aerobic. Cylindrical cells, 3-4 times as long as broad, united into 
long rod-like bodies, which may elongate into long, bent, and 
twisted filaments. Not self-motile. Endosporous. Germination 
takes place without the throwing off of any spore-membrane (com- 
pare Hay-bacillus p. 37 which resembles it). Contagion may take 
place both by introduction into wounds, and from the mucous 
membrane of the intestines or lungs, both by vegetative cells and 
by spores ; in intestinal anthrax, however, only by spores. The 
Bacillus multiplies as soon as it has entered the blood, and the 
anthrax disease commences. The Bacilli not only give off poison, 



but also deprive the blood of its oxygen. Vegetative cells only 
occur in living animals. This species is a, facultative parasite which 
in the first stage is a saprophyte, and only in this condition forms 

Bacillus tuberc^llosis produces tuberculosis in human beings, also 
in domestic animals (perlsucht). It is a distinct parasite, but may 
also live saprophytically. It is rod-formed, often slightly bent, 
and is recognised principally by its action with stains (when 
stained with an alkaline solution of methyl-blue or carbolic fuch- 

PI --*, 

=8 -\\ 

^ \ 

. i -'= o. iSk 

^-B : : S- 

M: -I 

FIG. 32. Anthrax bacillus (Bacil- 
lus anthracis) with red (b) and white 
(a) blood-corpuscles. 

FIG. 33. Anthrax bacillus. The formation of 
the spores ; magnified 450 times. 

sin, it retains the colour for a long time even in solutions of mineral 
acids, in contrast with the majority of well-known Bacteria): it 
probably forms spores which are able to resist heat, dryness, etc. 

Bacillus lepne produces leprosy ; Bacillus mallei produces glanders ; Bacillus 
tetani, tetanus (the tetanus bacillus is very common in soil ; anaerobic) ; Bacillus 
diphtheria, diphtheria ; Bacillus typhosus, typhoid fever, etc. 

Pathogenic Spiral Bacteria. Spirochwte oberm&ieri (Fig. 
24) produces intermittent fever (febris recurrens) ; it makes its 
appearance in the blood during the attacks of fever, but it is not 
to be found during intervals when there is no fever. Obligate 

Spirillum cholerce asiaticcB (Microspira comma) without doubt 
produces Asiatic cholera ; an exceedingly motile spirillum, which 
is also found in short, bent rods (known as the "Comma-bacillus"), 



it lives in the intestines of those attacked by the disease, and 
gives off a sti*ong poison which enters the body. It is easily 
ciiltivated as a saprophyte. 

A great many circumstances seem to show that a number of 
other infectious diseases (syphilis, small-pox, scarlet-fever, measles, 
yellow-fever, etc.) owe their origen to parasitic Bacteria, but this 
has not been proved with certainty in all cases. 

It has been possible by means of special cultivations (ample 
supply of oxygen, high temperature, antiseptic materials) to pro- 
duce from the parasitic Bacteria described above (e.g. the fowl-cholera 
and the anthrax Bacteria) physiological varieties which are distinct 
from those appearing in nature and possess a less degree of 
" virulence," i.e. produce fever and less dangerous symptoms in 
those animals which are inoculated with them. The production 
of such physiological varieties has come to be of great practical 
importance from the fact that they are used as vaccines, i.e. these 
harmless species produce in the 
animals inoculated with them 
immunity from the malignant in- 
fectious Bacteria from which they 
were derived. This immunity is 
effected by the change of the 
products of one or more of the 
Bacteria, but we do not yet know 
anything about the way in which 
they act on the animal organism. 
The white blood corpuscles, ac- 
cording to the Metschnikoff, play 
the part of " Phagocytes " by 
absorbing and destroying the less virulent Bacteria which have 
entered the blood, and by so doing they are gradually enabled to 
overcome those of a more virulent nature. 

Class 5. Conjugatae. 

The Algae belonging to this class have chlorophyll, and pyre- 
noids round which starch is formed. The cells divide only in one 
direction, they live solitarily, or united to form filaments which 
generally float freely (seldom attached). Swarm-cells are want- 
ing 1 . The fertilisation is isogamous (conjugation) and takes place by 
means of apJanogametes. The zygote, after a period of rest, 
produces, immediately on germination, one or more new vegetative 

FIG. 31. a and i> The same blood-cell 
of a Frog: a in the act of engulfing 
an anthrax-bacillus ; b after an interval 
of a few minutes when the bacillus has 
been ab? orbed. 



individuals ; sometimes akinetes or aplanospores are formed in 
addition. They only occur in fresh or slightly brackish water. 

Order 1. Desmidiaceae. The cells generally present markings 
on the outer wall, and are mostly divided into two symmetrical 
halves by a constriction in the middle, or there is at least a 
symmetrical division of the protoplasmic cell-contents. The cell- 
wall consists nearly always of two layers, the one overlapping 
the other (Fig. 35 0). The cells either live solitarily or are united 
into unbranched filaments. The mass of protoplasm formed by 
the fusion of the two conjugating cells becomes the zygote, which 
on germination produces one (or after division 2, 4 or 8) new 
vegetative individual. The chromatophores are either star-, plate-, 
or band-shaped, and regularly arranged round the long axis of the 


FIG. 35. A Cell of Gi/wiiozy.oa brebistonii, external view showing the distribution of the 
pores. B A portion of the membrane of Slaurastrum licorne with pores containing proto- 
plasmic projection?. Cell-wall of Hyalotheca inucosa during cell-division ; the central 
part, being already formed, shows the connection with the divisional wall. 

The Desmidiaceee are not able to swim independently, many 
species, however, show movements of different kinds by rising and 
sliding forward on the substratum. These movements, which are 
partly dependent upon, and partly independent of light and the 
force of gravitation, are connected with the protrusion of a mu- 
cilaginous stalk. The mucilage, which sometimes surrounds the 
whole individual, may acquire a prismatic structure, it is secreted 
by the protoplasmic threads which project through certain pores 
definitely situated in the walls (Fig. 35 A, B). 

VEGETATIVE MULTIPLICATION takes places by division. A good 
example of this is found in Cosmarium botrytis (Fig. 36 A-D). 
The nucleus and chromatophores divide, and simultaneously the 
central indentation becomes deeper, the outer wall is then ruptured 
making a circular aperture through which the inner wall protrudes 
forming a short, cylindrical canal between the two halves to which 
it is attached (Fig. 36 C). After elongation the canal is divided 
by a central transverse wall, which commences as a ring round its 


inner surface and gradually forms a complete septum. The divid- 
ing wall gradually splits, and the two individuals separate from 
each other, each one having an old and a new half. The two 

FIG. 36. Cosmo rinm botrytis. A-D Different stages of cell-division. 

daughter-cells bulge out, receive a supply of contents from the 
parent-cells, and gradually attain their mature size and develop- 
ment (Fig. 36-B-D). Exceptions to this occur in some forms. 

CONJUGATION takes place in the simplest way in Mesotcenium, where 
the two conjugating cells unite by a short tube (conjugation-canal), 
which is not developed at any particular point. The aplanogametes 
merge together after the dissolution of the dividing wall, like two 
drops of water, almost without any trace of preceding contraction, 
so that the cell-wall of the zygote generally lies in close contact 
with the conjugating cells. The conjugating cells in the others 
lie either transversely (e.g. Cosmarium, Fig. 37 d ; Staurastrum, 
etc.), or parallel to one another (e.g. Penium, Closterium, etc.), and 

FIG. 37. Cosmarium menegliinii : a-c same individual seen from the side, from the end, 
and from the edge ; d-f stages of conjugation ; g-i germination of the zygote. 

emit a short conjugation-canal (Fig. 37 d) from the centre of that 
side of each cell which is turned towards the other one. These 
canals touch, become spherical, and on the absorption of the divid- 
ing wall the aplanogametes coalesce in the swollen conjugation- 
canal (Fig. 37 c), which is often sin-rounded by a mucilaginous 
envelope. The zygote, which is often spherical, is surrounded by 
a thick cell-Avail, consisting of three layers ; the outermost of these 


sometimes bears thorn-like projections, which in some species 
are simple (Fig. 37/), in others branched or variously marked; 
in some, however, it remains always smooth (e.g. Tetmemorus, 
Desmidium*). Deviation from this mode of conjugation may occur 
within certain genera (e.g. Closterium, Penium). Upon germina- 
tion the contents of the zygote emerge, surrounded by the inner- 
most layers of the wall (Fig. 37 g, h\ and generally divide into 
two parts which develope into two new individuals, placed trans- 
versely to each other (Fig. 37 i) ; these may have a somewhat more 
simple marking than is generally possessed by the species. 

FIG. 38. Destnidiacese. A Closterium moniliferum ; Ji Pennon crastiu&culum ; C 
Micrasterias truncata .(front and end view) ; D Euastruin, elegans ; E Stauraktrum muticum 
(end view). 

The most frequent genera are : 

A. Solitary cells : MESOT^NIUM, PENIUM (Fig. 38 ?), CYLINDKOCVSTIS, 
EUASTRUM (Fig. 38 D), MICRASTEEIAS (Fig. 38 C), COSMAHIUM (Fig. 36, 37), 

B. Cells united into filaments : SPH.EROZOSMA. DESMIDICM, HYALOTHECA, 

Order 2. Zygnemaceae. Cell-wall without markings. The 
cells are cylindrical, not constricted in the centre, and (generally) 
united into simple, unbranched filaments. The whole contents of 
the conjugating cells take part in the formation of the zygote, 
which on germination grows out directly into a new filament. 

Spirogyra is easily recognised by its spiral chlorophyll band ; 
Zy/jnema has two star-like chromatophores in each cell (Fig. 40) ; 
both these genera are very common Alga? in ponds and ditches. 



The conjugation among the Zygnemacete takes place in the 
following manner : the cells of two filaments, lying side by side, or 

A n 

FIG. 39. Spirogyra longata. A At the commencement of conjugation, the conjugation- 
canals begin to protrude at a and touch one another at J> ; the spiral chlorophyll band and 
cell-nuclei (fc) are shown. B A more advanced stage of conjugation; a, a' the rounded 
female and male aplanogametes : in b' the male aplanogamete is going over to and 
uniting with the female aplanogamete (b). 

two cells, the one being situated above the other in the same fila- 
ment (Fig. 41), push out small protuberances opposite each other 
(Fig. 39 A, a, b) ; these finally meet, and the dividing wall is ab- 
sorbed so that a tube is formed connectin one cell with the other ; 


FIG. 40. A cell of Zygnema. S Pyrenoid. 

FIG. 41. Zygnema insigne, with zygote. 

the protoplasmic contents round off, and the whole of these contents 
of one of the cells glides through the conjugation-tube and coalesces 

with that of the 
other (Fig. 39 5), 
the aggregate mass 
then rounds off, sur- 
rounds itself with 

FIG. 42. Germinating zygote of Spirogyra jugalis -. the a cell-Wall, and be- 

young plant is still unicellular; the end which is still in comes a ZV^ote A 
the wall of the zygote is elongated and root-like; the 

chromatophore divides and forms the spiral band. distinct difference 


may be found between the cells in the two filaments, those in the 
one whose protoplasmic contents pass over being cylindrical, while 
those of the recipient one are more barrel-shaped, and of a larger 
diameter. The former may be regarded as a male, the latter as 
a female plant. The zygote germinates after a period of rest, and 
grows out into a new filament (Fig. 42). 

Order 3. Mesocarpaceae. The cell-walls are glabrous, uncon- 
stricted in the centre, and united into simple unbranched filaments. 
The chromatophore consists of an axial chlorophyll-plate, with 
several pyrenoids. The zygote is formed by the coalescence of two 
cells (Fig. 43) (sometimes three or four), but the whole proto- 
plasmic contents of the cells do not take part in this process, a 
portion always remaining behind ; the aplanogametes coalesce in 
the conjugation-canal. The zygote thus formed appears incapable 
of germination until after 3-5 divisions. Of the cells so formed, 

FIG. 43. Nowjeotia calcarea. Cells showing various modes of conjugation : at m tripar- 
tition ; pg quadripartition ; s quinquipartition of the zygote. 

only one is fertile, the sterile cells, according to Pringsheim, con- 
stituting a rudimentary sporocarp. The germinating cells grow 
out into a new filament. In this order, conjugation has been ob- 
served between two cells of the same filament. The Mesocarpacese 
thrive best in water which contains lime. 

Class 6. Chlorophyceae (Green Algae). 

These Algse are coloured green by chlorophyll, seldom in com- 
bination with other colouring matter, and then especially with red. 
The product of assimilation is frequently starch, which generally 
accumulates round certain specially formed portions of protoplasm 
termed pyrenoids. The thallus is uni- or multi-cellular; in the 
higher forms (certain Siphonea?) the organs of vegetation attain 
differentiation into stem and leaf. The asexual reproduction takes 
place in various ways ; the sexual reproduction is effected by con- 
jugation of motile gametes, or by oogamous fertilisation. The 


swarm-cells (zoospores, gametes, and spermatozoids) are con- 
structed symetrically, and have true protoplasmic cilia, these 
generally being attached to the front end of the swarm- cells. 
Most of these Alga? live in water (fresh or salt); some are found 
upon damp soil, stones, or tree- stems, and some live enclosed in 
other plants. 

The Class is divided into three families : 

1. PROTOCOCCOIDEJ; : Volvocaceae, Tetrasporacea?, Chloro- 
sphasraceas, Pleurococcaceaa, Protococcaceas, Hydrodictyaceae. 

2. CONFERVOIDE^E : Ulvacea?, Ulothricaceae, Cha?tophoracea?, 
Mycoideacea3, Cylindrocapsaceaa, (Edogoniacege, Coleochaetacea-, 
Cladophoraceas, Gomontiacea?, Sphn?ropleacea3. 

3. SIPHONED : Botrydiacea?, Bryopsidacea?, Derbesiaceae, Vau- 
cheriacea?, Pbyllosiphonacea 3 , Caulerpacese, Codiaeese, Valoniaceae, 

Family 1. Protococcoideae. 

The Alga 1 which belong to this group are uni- or multi-cellular 
with the cells more or less firmly connected, sometimes in a 
definite, sometimes in an indefinite form (Fig. 47). Colonies are 
formed either by division or by small unicellular individuals be- 
coming united in a definite manner ; the colonies formed in this 
latter way are termed C&nobia. Apical cells and branching are 
absent. Multiplication by division ; asexual reproduction by zoo- 
spores, rarely by akinetes. Sexual reproduction may be wanting, 
or it takes place by isogamous, rarely by oogamous fertilisation. 

Some are attached by means of a stalk to other objects (Chara- 
cium, Fig. 49), others occur as " Endophytes " in the tissues of 
certain Mosses or Phanerogams, e.g. Chlorochytrium lemncu, in Lemna 
trisitlca ; Endosphcera, in the leaves of Potamogeton, Mentha aquatica, 
and Peplis portula ; Phyllobium, in the leaves of Lynmachia num- 
inularia, A/uga, Chlora, and species of Grasses ; Scotinosphcera in 
the leaves of Hypnum and Lemna trisulca ; the majority, however, 
live free in water and in damp places. Many species which were 
formerly considered to belong to this family have been proved to 
be higher Alga? in stages of development. 

Order 1. Volvocaceae. The individuals in this order are either 
uni- or multi-cellular, and during the essential part of their life 
are free-swimming organisms. They are generally encased in a 
mucilaginous envelope, through which 2-6 cilia project from every 


cell. The vegetative reproduction takes place by the division of 
all, or a few, of the cells of the individual ; in some a palmella-stage 
is found in addition. The sexual reproduction takes place by 
isogamous or oogamous fertilisation. 

The Volvocacese may be considered to include the original forms of the Chloro- 
phycete, because, among other reasons, the motile stage is here the most promi- 
nent ; they also form the connecting link between the animal Flagellata, and 
forms intermediate to the Syngeneticcc may perhaps be found amongst them. 
Three series of green Algee may be supposed to have taken their origin from the 
Volvocacea? : CONJUGATED (Desmidiacece) which have lost the swarming stage, but 
whose conjugation is the nearest to the fertilisation in Chlannjdomonas pulvis- 
cuhis : the PKOTOCOCCACE.S: in which the vegetative divisions have disappeared, 
while the swarming stage continues to be present, though of shorter duration ; 
and TETRASPORACE^E, in which the vegetative divisions are more prominent, 
whilst the swarming stage is less so. 

A. UNICELLULAR INDIVIDUALS. The principle genera are : Chla- 
mydomonas, Sphcerella, Pliacotns. Sphterella nivalis is the Alga 
which produces the phenomenon of " Red Snow," well known en 
high mountains and on ice and snow fields in the polar regions. 
The red colouring matter which appears in this and other green 
Alga?, especially in the resting cells, is produced by the alteration 
of the chlorophyll. 

Phacotus lenticidaris has an outer covering incrusted with lime, 
which, at death, or after division, opens out into two halves. 
Species may be found among Chlamydomonas, in which conjuga- 
tion takes place between gametes of similar size without cell-wall, 

but in C. pulvisculus conjugation 
takes place between male and 
female aplanogametes which are 
surrounded by a mucilaginous 


The most important genera are 
Gonium, Stephanosplicera, Pandor- 
inn, Eudorina, Volvox* Gonium 
has 4 or 16 cells arranged in a 
definite pattern in a flat plate 

Fig. 44. Gonium pectornle. /T -,. ,. x -^ , . -.-r-,. . < 

(Fig. 44). Pandonna (Fig. 45), 

has 16 cells arranged in a sphere (Fig. 45 A). The vegetative 
reproduction takes place in this way : each cell, after having 
rounded off, and after the withdrawal of the cilia, divides itself 



into 16 new ones (Fig. 45 B), each forming a new individual, 
which soon grows to the size of the mother-individual. It was 
in this Alga that the conjugation of self-motile gametes was 
first discovered by Pringsheim, 1869. When conjugation is 
about to take place, each cell divides into sixteen, as in vegeta- 
tive reproduction, but the 16 x 16 cells all separate from one 
another (Fig. 45 C, female gametes, and D, male gametes), and 

FIG. 45. Pandorina worum. 

swarm solitarily in the water. The male are, most frequently, 
smaller than the female, but otherwise they are exactly alike ; 
they are more or less pear-shaped, with a colourless anterior end, 
2 cilia, a red " eye-spot," etc. After swarming for some time they 
approach each other, two and two, generally a large and a smaller 
one, and come into contact at their colourless end ; in a few 
moments they coalesce and become one cell (Fig. 45 E, F), this 

W. B. E 



has at first a large colourless anterior end, 4 cilia, and 2 " eye- 
spots" (Fig. 45 G), but these soon disappear and the cell becomes 
uniformly dark-green and spherical, and surrounds itself with a 
thick cell-wall, losing at the same time its power of motion : the 
zygote (Fig. 45 H) is formed, and becomes later on a deep red 
colour. On the germination of the zygote, the protoplasmic cell- 
contents burst open the wall (Fig. 45 J), and emerge as a large 
swarmspore (Fig. 45 K) which divides into 16 cells, and the first 
small individual is formed (Fig. 45 i, M). 

Eudorina is like Pandorina in structure, but stands somewhat 

FIG. 46. Volvox gfobator, sexual individual : a antheridia which have formed sperniato- 
zoids ; b oogonia. 

higher, since the contrast between the conjugating sexual cells is 
greater, the female one being a motionless oosphere. 

The highest stage of development is found in Volvox (Fig. 46). 
The cells are here arranged on the circumference of a sphere, and 
enclose a cavity filled with mucilage. The number of these cells 
may vary from 200-22,000, of which the majority are vegetative 
and not reproductive, but some become large, motionless oospheres 
(Fig. 46 b) ; others, which may appear as solitary individuals, 
divide and form disc- shaped masses of from 8-256 small spermato- 



zoids (Fig. 46 a). After the oosphere has been fertilised by these, 
the oospore surrounds itself by a thick, sometimes thorny cell- 
wall, and on germination becomes a new individual of few cells. 
A few cells conspicuous by their larger size may be found (19, but 
generally 8) in certain individuals, and these provide the vegeta- 
tive reproduction, each forming by division a new individual. 

Order 2. Tetrasporaceae reproduce both by vegetative divisions and 
swarmspores, some have also gamete-conjugation. The principal genera are : 
Tetraspora, Apioci/stis, Dactylococcus, Dictyuspharium, Chloranyium. 

Order 3. Chlorosphaeraceae. Chlurosphara. 

Order 4. Pleurococcaceae. In this order the swarm-stages 


and sexual reproduction are entirely absent. Vegetative repro- 
duction by division. The principal genera are : Pleurococcus (Fig. 
47), Scenedesmus (Fig. 48), . Raphidium, Oocystis, ScMzochlamys, 
Crucigenia, Selenastrum. Pleurococcus vulgaris (Fig. 47) is one of 
the most common Algae throughout the world, occurring as o'reen 
coverings on tree-stems, and damp walls, and it is one of the most 
common lichen-gomdia. 


FIG. -17. Pleurococcus vulgaris. 

FIG. 48. ScciictlcsiiiHS quadricauda. 

Order 5. Protococcaceae. The cells are motionless, free or 
affixed on a stalk (e.g. Characium, Fig. 49), either separate or 
loosely bound to one another ; they never form multicellular 
individuals. Multiplication by division is 
nearly always wanting. Reproduction takes 
place by swarmspores, which have 1 or 2 cilia, 
and sexual reproduction in some by gamete- 
conjugation. The principal genera are : Chlo- 
rococcum, Ghlorochytfium, Chlorocystis, Scotino- 
sphc&ra, Endosplicera, PhylJobium, Characiurn, 
Ophwcytium, Sciadium. 

Order 6. Hydrodictyacese. The indi- 
viduals are unicellular but several unite after 
the zoospore-stage into definitely formed 
families (coenobia). Ordinary vegetative division is wanting, but 

FIG. 49. Characiwm 

strictum. A The cell-son- 
tents have divided into 
many swarrnspores. B 
Swarmspores escaping. 


asexual reproduction takes place by zoospores (or by motionless 
cells without cilia), which unite and form a family similar to the 
mother-family, inside the mother-cell, or in a mucilaginous enve- 
lope. Where sexual reproduction is found it takes place by 
gamete-conjugation. The principal genera are : Pediastrum (Fig. 
50), Cwlastrum, Hydrodictyon (Fig. 51). 

The coanobiuin of Hydrodictyon reticulatum (Water-net) is 
formed of a large number of cells which are cylindrical, and 
attached to one another by the ends (Fig. 51). The asexual 
reproduction takes place by zoospores, which are formed in large 
numbers (7,000-20,000) in each mother-cell, within which they 
move about for a time, and then come to rest and arrange them- 
selves into a new net (Fig. 51 A) which is set free by the dissolu- 
tion of the wall of the mother-cell, grows, and becomes a new 


FIG. 50. Pediastrum agperwm. FIG. 51. Hydrodictyon reticulatum. 

A A cell where the zoospores are on the 
point of arranging themselves to form a 
net. B A cell with gametes svyarming out. 

coenobium. The sexual reproduction takes place by gamete-conju- 
gation. The gametes are formed in the same manner as the zoo- 
spores, but in larger numbers (30,000-100,000), and swarm out of 
the mother-cell (Fig. 51 B). The zygote forms, on germination, 
2-5 large zoospores, each with one or two cilia, these generally 
swarm about for a time, and after a period of rest become irregular 
thorny bodies (polyhedra) ; their contents again divide into zoo- 
spores, the thorny external coating of the polyhedra is cast off, 
and the zoospores, surrounded by the dilated internal coating, 
unite to form a small family, which produces several others in the 
manner described. 


Family 2. Confervoideae. 

The individuals are always multicellular, the cells firmly 
bound together and united into unbranched or branched filaments, 
expansions, or masses of cells which grow by intercallary divisions 
or have apical growth. In the first seven orders the cells are 
uninuclear, but the cells of the remaining three orders contain 
several nuclei. Asexual reproduction by zoospores, akinetes or 
aplanospores. Sexual reproduction by isogamous or oogamous 

The Confervoidea?, through the Ulvaceaj, are connected with the Tetra- 
sporacefe, and from the Coleochccttrcefe, which is the most highly developed 
order, there are the best reasons for supposing that the Mosses have taken 
their origin. The Clndophoracea: show the nearest approach to the Sipliontie. 

Order 1. Ulvaceae. The thallus consists of one or two layers, 
of parenchymatous cells, connected together to form either a flat 
membrane (JMonnstroma, Ulva) or a hollow tube (Enteromorplia), 
and may be either simple, lobed, or branched. Reproduction 
takes place by detached portions of the thallus ; or asexually by 
zoospores or akinetes. Gamete conjugation is known to take 
place in some members of this order, the zygote germinating 
without any resting-stage. The majority are found in salt or 
brackish water. 

Order 2. Ulothricaceae. The thallus consists normally of a 
simple unbranched filament (sometimes a small expansion con- 


FIG. 52. Ulothrix zonnta: a portion of a filament with zoospores, which are formed 
two in each cell (zoosporangium); the dark spots are the red "eye-spots"; 1, 2, 3, 4, 
denote successive stages in the development of the zoospores ; b a single zoospore, v 
the pulsating vacuole ; c portion of a filament with gametes, sixteen are produced in each 
gnmetangium ; d free gametes, solitary or in the act of conjugation ; e the conjugation is 
completed, and the formed zygote has assumed the 

sisting of one layer of cells is formed, as in Schizomeris and 
Prasiola which were formerly described as separate genera). 
Asexual reproduction takes place by means of zoospores (with 


1, 2, or 4 cilia), akinetes or aplanospores ; the last named may 
germinate immediately, or only after a period of rest. Sexual re- 
production takes place by the conjugation of gametes of about the 
same size, each having two cilia (Fig. 52 d). The zygote of 
Ulothrix, on germination, produces a brood of zoospores which 
swarm for a time and then elongate to become 7/o^/m',-filaments 
(alternation of generations). The gametes may also germinate 
without conjugation in the same manner as the zoospores. The 
principal genera are : Ulothrix, Hormidium, Conferva, Micro- 
spora. Ulothrix zonata is very common in running fresh water. 
Nearly all the species of Hormidium occur on damp soil, tree- 
stems and stones. 

Order 3. Chaetophoraceae. The thallus consists of a single, 
branched, erect or creeping filament of cells, often surrounded by 
mucilage. The cells have only one nucleus. Asexual reproduc- 
tion by zoospores with 2 or 4 cilia, by akinetes, or aplanospores. 
In many, conjugation between gametes with 2 cilia may be found. 
They approach on one side, Ulothricaceae, and on the other, My- 
coideacea?. The principal genera are : Siigeoclonium, Draparnahlia, 
Cluvtophora, Enloderma, Aphanochtvte, Herposteiron, Phceuthamnion, 
Cltlorotylium, Trichophihis, Gongrosira, TrentepohJia. Most of the 
species of Trentepoklia are coloured red by the presence of a red 
colouring material, which occurs in addition to the chlorophyll. 
They are aerial Algae which live on stones (T. julithus, "violet 
stone," so named on account of its violet-like odour in rainy 
weather), on bark and old wood (T. umbrina'), or on damp rocks 
(T. aurea). Trichophilus icelckeri lives in the hair of Bradypus. 

Order 4. Mycoideaceae. The thallus is discoid, consisting of one or more 
cell-layers, and is always attached. Asexual reproduction by zoospores with 2 
or 4 cilia. Sexual reproduction in some species by the conjugation of gametes 
with 2 cilia. This order forms the connecting link betweeu Cluctop]ior(tct<c 
and Coleochcetacea. The species occur in fresh water (Cluetopeltis) as well as 
iu salt (Pringtsheimia), on the carapace of tortoises (DermatOphyton = Epiclem- 
mydia), or endophytic between the cuticle and the epidermal cells of the 
leaves of tropical plants, destroying the leaf-tissue (Mycoidea). 

Order 5. Cylindrocapsaceae. The thallus consists of a 
simple (rarely, in parts, formed of many rows) unbranched 
filament, attached in the young condition, which has short cells 
with a single nucleus, and is enveloped in a thick envelope with 
a laminated structure. Asexual reproduction by zoospores with 
2 cilia, which are formed 1, 2, or 4 in each vegetative cell. The 



antheridia are produced by a single cell, or a group of cells, in a 
filament, dividing several times without increasing in size. Two 
egg-shaped spermatozoids, each with 2 cilia (Fig. 53 D), are 
formed in each antheridium, and escape through an aperture in 
the side ; in the first stages they are enclosed in a bladder-like 
membrane (Fig. 
53 B, C). Other 
cells of the fila- 
ment swell out 
and form oogonia 
(Fig. 53-4), which 
resemble those 
of CEdogonium. 
After fertilisa- 
tion, the oospore 
surrounds itself 
with a thick wall, 
and assumes a 
reddish colour. 
The germination 
is unknown. 
The unfertilised 
oospheres remain 
green, divide often 
into 2-4 daughter- 
cells, and grow 
into new fila- 

This order, 
which 'only in- 
cludes one genus, Cylindrocapsa, forms the connecting link be- 
tween Ulothricacece and (Edogoniacece. The few species (4) occur 
only in fresh water. 

Order 6. CEdogoniaceae. The thallus consists of branched 
(Bulbochcete) or unbranched (CEdogonium) filaments, attached in 
the early stages. The cells may be longer or shorter, and have 
one nucleus. Asexual reproduction by zoospores, which have a 
chaplet of cilia round the base of the colourless end (Fig. 6 a). 
Sexual reproduction takes place by oogamous fertilisation. On 
the germination of the oospore, 4 zoospores are formed (Fig. 54 F). 
They occur only in fresh or slightly brackish water. The division 



FIG. SS.Cylindroeapsainvoluta. A Oogonium withoosphpre 
(o) surrounded by spermatozoids (s). B Two antheridia, each 
with two sperraatozoids. C Spermatozoids surrounded by 
their bladder-like membrane. D Free spermatozoid. 



of the cells takes place in quite a peculiar and unusual manner. 
At the upper end of the cell which is about to divide, a ring-shaped 
thickening of soft cellulose is formed transversely round the wall ; 
the cell-nucleus of the mother-cell and the protoplasm then divide 
by a transverse wall into two portions of similar size, and the 

cell-wall bursts trans- 
versely along the cen- 
tral line of the thick- 
ened ring. The cell- 
wall thus divides into 
two parts the upper 
one short, the "cap," 
and the lower one 
much longer, the 
"sheath." The por- 
tions of the original 
cell-wall now separate 
from each other, the 
cellulose ring extend- 
ing, and supplying 
an additional length 
of cell-wall between 
them. The cap and 
sheath will project a 
little in front of the 
piece thus inserted. 
The dividing wall be- 
tween the two new 
cells is formed near to 
the uppermost edge 
of the sheath, and 
gradually becomes 
thicker and firmer. 
The inserted piece of 
wall forms the larger 
part of the wall of the 
upper cell : the re- 
mainder is formed by 

FIG. 51. A (Edogonium eilintum. A Female plant with 
three oogonia (03) and dwarf-males (m). B An oogonium 
with spermatozoid (z) seen entering the oosphere (n) hav- 
ing passed through an aperture near the summit of the 
oogonium; m dwarf-male. C Ripe oospore. D (Edogo- 
nium gcmeUiparum. F Portion of a male filament from 
which spermatozoids (z) are emerging. E Portion of 
filament of Btdboc)i<rfc ; the upper oogonium still en- 
closes the oospore, in the central one the oospore is es- 
caping while the lower one is empty. F Four zoospores 
developed from an oospore. G Zoospore germinating. 

the cap. This mode 
of division is repeated exactly in the same way, and new caps are 
formed close below the first one, one for every division. 



Fertilisation takes place in the following way. The oogonium. 
is a large ellipsoidal, swollen cell (og, in Fig. 54 A), whose contents 
are rounded off into an oosphere with a colourless receptive-spot 
(see B} ; an aperture is formed in the wall of the oogonium, through 
which the spermatozoids are enabled to enter (_B). The sper- 
matozoids are pi^oduced either directly, as in D (in pairs), in basal 
cells of the filament, or indirectly. In the latter case a swarm- 
spore (androspore) is formed which comes to rest, attaches itself 

FIG. 65. CoJeocJusfe pulrinatn. A A portion of a thallus with organs of reproduction; 
a oogonium before, b after fertilisation ; can antheridium, closed; d open, with emerg- 
ing spermatozoid. -B Ripe oogonium, with envelope. C Germination of the oospore. T> 
Zoospore. E Spermatozoid. 

to an oogonium, germinates, and gives rise to a filament of a very 
few cells dwarf-male (A, B, ?). The spermatozoids are formed 
in the upper cell of the dwarf-male (i), and are set free by the 
summit of the antheridium lifting off like a lid. On the germina- 
tion of the oospore (C), which takes place in the following spring, 
4 zoospores are produced (F) (i.e. the sexual generation) ; these 
swarm about for a time, and ultimately grow into new filaments. 

Order 7. Coleochsetacese. The thallus is always attached, 
and of a disc- or cushion-shape, formed by the dichotomous 
branching of filaments of cells united in a pseudo-parenchy- 
matous manner. Each cell has only one nucleus. Asexual re- 
production by zoospores with 2 cilia (Fig. 55 D), which may 
arise in all the cells. Sexual reproduction by oogamous fertili- 
sation. The spermatozoids resemble the swarmspores, but are 


smaller (E), and originate singly (in the species figured) in small 
conical cells (c, d in A\ The oogonia are developed at the 
extremities of certain branches : they are bottle-shaped cells with 
very long and thin necks (trichogyne), open at the end (a in A); 
at the base of each oogonium is a spherical oosphere. The sper- 
matozoids reach the oosphere through the trichogyne, or through 
an aperture in the wall when the trichogyne is absent, and fertili- 
sation having taken place, the oogonium becomes surrounded by a 
cell-layer (envelope), which grows out from the cells near its base 
(b in A), and in this way a kind of fruit is formed (B) (spermocarp, 

cystocarp) . 

The oospore, next spring, divides and forms a parenchymatous 
tissue (homologous with the Moss-sporophyte) ; this bursts open 
the envelope (C), and a zoospore (homologous with the spores of 
the Moss-capsule) arises in each of the cells, and produces a new 
Goleochcete. We have then, in this case, a still more distinct 
alternation of generations than in GEdogonium. Only one genus, 
Coleochcete, is known, but it contains several species, all living in 
fresh water. 

Order 8. Cladophoraceae. This order is probably derived 

from the Ulothricacea?. The thallus consists of a single, un- 

branched or branched filament, generally with an apical cell. 

The cells have each 2 or more nuclei. Asexual reproduction by 

zoospores with 2 or 4 cilia, and by akinetes. Conjugation of 

gametes with 2 cilia is found in some genera. They occur in salt 

as well as in fresh water. The principal genera are: Urospora, 

Chcetomorpha, Ithizoclonium, Cladophora; of the last named genus 

the species C. lanosa and C. rupestris are common in salt water ; 

C. fracta and C. glomerata in fresh water. 

Order 9. Gomontiace*. Gomontia polyrrhiza, the only species hitherto 
known, is found on old calcareous shells of certain salt water Molluscs. 

Order 10. Sphceropleaceae. The thallus consists of free, un- 
branched filaments, with very elongated multinuclear cells. The 
ve-etative cells form no zoospores. Sexual reproduction by 
oogamous fertilisation (see page 13, Fig. 10 B). The oospore has 
a thick wall (Fig. 10 D) studded with warts, and assumes a 
colour resembling red lead. It germinates only in the following 
spring, and produces 1-8 zoospores, each with 2 cilia (Fig. 10 E), 
which grow into new filaments. Only one species, Sphceroplea 
unnulina, is known. 


Family 3. Siphoneae. 

The thallus has apical growth, and in the vegetative condition 
consists generally of one single (in the Valoniacese most frequently 
of more) multinuclear cell, which may be much branched, and 
whose separate parts in the higher forms (e.g. Bryopsi*, Fig. 57 ; 
Caulerpa, Fig. 59, etc.) may be differentiated to perform the 
various physiological functions (as root, stem and leaf). Vegeta- 
tive multiplication by detached portions of the thallus (gemmae) ; 
asexual reproduction by zoospores, akinetes, or aplanospores. 
Sexual reproduction by gamete-conjugation, rarely by oogamous 
fertilisation. The zygote or oospore germinates as a rule without 
any resting-stage. 

Most of the Siphonene occur in salt water or on damp soil. 

FIG. 56. Botrydium granulatum : a an entire plant forming 1 swarmspores ; b swarm- 
spores; c an individual with gametangia ; d, gamete; e,f,g conjugation; h zygote 
seen from above ; i the same in a lateral view. 

Many (e.g. Dasycladact'W) are very much incrusted with lime, and 
occur, in the fossilized condition, in the deposits from the Cretaceous 
period to the present time. The Siphonete are connected by their 
lowest forms (Botrydiacew or Valonia) with the Protococcacea?, but 
show also, through the Valoniacese, points of relationship to the 

Order 1. Botrydiaceae. The thallus in the vegetative condi- 
tion is unicellular, club-shaped, with a small single (Codiolwrn) or 
repeatedly dichotomously branched system of colourless rhizoids 
(Botrydium, Fig. 56 a), by which it is attached to objects immersed 
in salt water (Codtolum) or to damp clay soil (-Botrydium). 
Asexual reproduction by zoospores with one (Botrydium) or two 



cilia, and by aplauospores. The sexual reproduction is only 
known in Botrydium, and takes place in the following manner: 
in the part of the thallus which is ahove ground and in an active 
vegetative condition, several round cells (Fig. 56 c) are formed, 
which may be green or red according as they grow under water, 
or exposed to the strong light of the sun. These cells must be 
considered as "gametangia" as they produce many gametes (d) 
provided with two cilia. The zygote (h, i) formed by the conju- 
gation (e, /, g) may either germinate immediately, or become a 
thick-walled resting-cell of an irregular, angular form. 

Order 2. Bryopsidacese. The thallus in the vegetative condition is uni- 
cellular, and consists at the 
lower extremity of branched 
rhizoids. while the upper 
portion is prolonged into a 
stem-like structure of un- 
limited growth, producing, 
acropetally, branches and 
leaf-like structures. The 
latter have limited growth, 
and are separated by a cross 
wall from the stem, and be- 
come gametangia, or drop 
off. The gametes have two 
cilia, and are of two kinds : 
the female, which are green 
and large and the male, 
which are of brownish 
colour and smaller. Zoospores or anj- other method of asexual reproduction 
are unknown. Only one genus, Bryopsis, living in salt water. 

OrderS. Derbesiacese. Only one genus, Derbesia, living in saltwater. The 
zoospores, which are formed in a few lateral, swollen zoosporangia, possess one 
nucleus which has arisen through the coalescence of several, and they resemble 
the zoospores of ffidogonium by having a circle of cilia attached at the base 
of the colourless spot. 

Order 4. Vaucheriacese. The thallus consists, in the vegetative 
condition, of a single irregularly or dichotomously branched cell, 
without differentiation into stem or leaf ; root-like organs of attach- 
ment may however occur. Asexual reproduction by zoospores, 
which are formed singly in the extremity of a branch cut off by 
a transverse wall. They contain many nuclei, and bear small 
cilia situated in pairs, which give the appearance of a fine " pile " 
covering the whole or a great part of the surface. Akinetes, 

FIG. 57. Bryopsis plumosa. A the plant, natural size. 
K A portion (enlarged) which shows the growing point 
(w), and the leaves derived from it in acropetal succes- 



aplanospores, and phytoamosbae (naked masses of protoplasm, 
without cilia, which creep like an amoeba on a substratum) may 
occur under certain conditions. 

The sexual reproductive organs are formed on short lateral 
branches, and are separated from the vegetative cell (Fig. 58 A) by 
cell-walls. Numerous spermatozoids, each with two cilia, are 
developed in the coiled antheridium (A, b). The oogonium is a 
thick, egg-shaped, often oblique cell, with its protoplasm rounded 
into an oosphere, which has a hyaline " receptive-spot" (A, a) im- 
mediately beneath the aperture formed in the wall of the oogonium. 
A slimy mass, which serves to receive the spermatozoids, is formed 
in some species in this aperture. The spermatozoids when liber- 
ated swim towards and enter the oosphere, which then immedia- 

A B 

FIG. 58. Vawcheria sessilis. A Fertilisation ; b the antheridia ; a the oogonia ; a the 
receptive spot. B Oospore. 

tely surrounds itself with a thick cell-wall. The mature oospore 
(-B) contains a large quantity of oil. At germination the outer 
cell-wall bursts and a new plant is formed. There is only one 
genus, Vaucheria, with species living in salt as well as in fresh 
water and on damp soil. 

Order 5. Phyllosiphonaceze are parasites in the leaves and stalks of 

Order 6. Caulerpaceae. The thallus has distinct dift'erentation 
into root, stem and leaf-like members (Fig. 59) ; it is unicellular. 
Within the cell, strong, branched threads of cellulose extend 
from one side to the other serving as stays to support the 
thallus. Reproduction takes place by detached portions of the 
thallus; no other modes of reproduction are known. This order 
may most approximately be classed with the Bryopsidacece. The 
genus Caulerpa consists of more than seventy species which 
inhabit the tropical seas. 

Order 7. Codiaceae. The thallus has various forms, but with- 


out distinct differentiation in stem- or leaf-structures, sometimes 
(e.g. Halimeda) it is very much incrusted with lime. In the 
early stages it is unicellular (later, often multicellular), very 
much branched, with the branches, at any rate partly, so united 
or grown in amongst one another (Fig. 60) that an apparently 
pai-enchymatous cellular body is formed. Akinetes or aplanospores 
are wanting; zoospores (or 
gametes ?) may be developed in 
some species, however, in special 
swollen sporangia. Fertilisation 
similar to that in Bryopsis occurs 
perhaps in Codium. They are 
rill salt water forms. 

Order 8. Valoniaceae. The thal- 
lus is generally multicellular, without 
differentation into stem- or leaf-struc- 
tures, but the cells are sometimes 
united together and form a leaf-like 
reticulate expansion (e.g. Anadyomene). 
Zoospores are known in some, and 
they are then formed directly in the 
vegetative cells. In others (e.g. 
Valonia), a mass of protoplasm, which 
maybe separated through the damag- 
ing of a cell, can surround itself with 

FIG. 59. Cnulcrpa, prolifera (natural size). 

a cell-wall, and grow into a new plant. No other modes of reproduction are 
known. The most important genera are : Valonia, Siphonocladus, Chamadoris, 
Strucea, Microtiictyon, Anadyomene. They are all salt water forms. 

As already pointed out, the Valoniaceer occupy a somewhat central position 
among the Siphoneze, and present points of similarity and contrast with the 
liotrydiacfa; and the Bryopsidacetc through Valonia, with the Dasycladaceee 
through C)iama>doris, and also with the Cladophoracete through Siphonocladus, 
and Slruvea. 



Order 9. Dasycladaceae. The thallus consists of an axile 
longitudinal cell, destitute of transverse walls, attached at the 
base by root-like organs of attachment, and producing acropetally 
whorls of united, single or branched, leaf-like structures with 
limited growth. Asexual reproduction is wanting. Sexual re- 
production by conjugation of gametes which arise in separate, 
fertile leaves/either directly or from aplanospores, which develope 
into gametangia. The principal genera are : Acetabularia, Dasy- 
cladus. Neomeris, Cymopnlia. All marine. 

The curiously shaped 
Acetalndaria mediterra- 
nea grows gregariously 
on limestone rocks, and 
shells of mussels in the 
Mediterranean ; it re- 
sembles a minute um- 
brella with a small stem, 
sometimes as much as 
nine centimetres in 
height, and a shade 
which may be more than 
one centimetre in di- 
ameter. The cell-mem- 
brane is thick, and 
incrusted with carbonate 
and oxalate of lime. 
Only the lower, root- 
like part of the thallus, 
which penetrates the 
calcareous substratum- 
survives the winter, and 
may grow up into a new 

plant. The sterile leaves, which drop off early, are dichoto- 
mously branched and formed of cylindrical cells separated 
from each other by cross-walls, but they are not grown to- 
gether. The shade is formed by a circle of 70-100 club-shaped 
rays (fertile leaves) grown together, in each ray 40-80 aplano- 
spores are formed, which become liberated at the breaking of the 
shade, and later on are changed to gametangia (compare Botry- 
dium} which open by a lid and allow a large number of egg- 
shaped gametes with two cilia to escape. Gametes from various 

FIG. 60. Halimeda opuntia. Plant (natural size). 
B Part of a longitudinal section. 


gametangia conjugate with one another; the product of the conju- 
gation swarms about for some time, rounds off, and then surrounds 
itself with a cell-wall. The zygote germinates after a period of 
rest and then produces a sexual plant. The aplanospores (game- 
tangia) thus represent the sexual generation. 

Class 7. Characese. 

The thallus has a stem with nodes and internodes ; and whorls 
of leaves, on which may be developed the antheridia and oogonia, 
are borne at the nodes. Vegetative reproduction by bulbils and 
accessory shoots. Zoospores are wanting. The antheridia are 
spherical, and contain a number of filaments in which the spirally 
coiled spermatozoids, each with two cilia, are formed. The oogonium 
is situated terminally, and is at first naked, but becomes later 
on surrounded by an investment, and forms after fertilisation the 
so-called " fruit." The oospore, after a period of rest, germinates 
by producing a " proembryo," from which the young sexual plant as a lateral branch. The Characeaa are distinguished by the 
structure of their vegetative system as well as by the spirally- 
coiled spermatozoids, and stand as an isolated group among the 
Thallophytes, of which, however, the Siphonete appear to be their 
nearest relations. They were formerly, but wrongly, placed near 
the Mosses. The class contains only one order, the Characese. 

Order 1. Characeae. Alga? with a peculiar odour, often 
incrusted with lime, and of a brittle nature. They generally grow 
gregariously in large masses at the bottom of fresh and brackish 
water, and are from a few inches to more than a foot in height. 
The stem has long internodes which in Nitella are formed of one 
cylindrical cell; in Chara of a similar cell, but closely surrounded 
by a cortical layer of smaller ones. The protoplasm in contact 
with the cell- wall exhibits in a well-marked degree the movement 
of rotation (cyclosis), carrying the chlorophyll corpuscles along 
with it. The internodes are separated from each other by a layer 
of small cells (nodal cells) from which the leaves are produced. 
The leaves are borne in whorls of from 5-12 which regularly 
alternate with one another as in the higher verticillate plants ; a 
branch is borne in the axil of the first formed leaf of each whorl 
(Fig. Gl A, ). 

The leaves are constructed in the same manner as the stem ; 
they are divided into a series of joints, but have only a limited 


power of growth ; their terminal cell, too, is not enclosed by a 
cortex. Leaflets are borne at their nodes. The growth of the 
stem is unlimited, and proceeds by means of an apical cell (Fig. 
62 s). The apical cell divides into a segment-cell and a new 
apical cell. The segment-cell then divides by a transverse wall 
into two cells, one lying above the other ; the lower one, without 

FIG. 61. Charafragilis. A Portion of a plant, natural size. B Portio'i of a leaf I, with 
leaflets '-" ; a antht-ridium ; c oogonium. C A shield. Nitella Jlexilis. D Filament from 
antherkliuru with sperm itozoids. E Free spermatozoids. 

any further division, becomes one of the long, cylindrical, inter- 
nodal cells (Pig. 62 in), and the upper one (Fig. 62 n) divides 
by vertical walls to form the nodal cells. The cortical cells 
(Fig. 62 r) which surround the long internodal cells of Chara, are 
derived from the divisions of the nodal cells ; the cells covering 
the upper portion of an internodal cell being derived from the 

W. B. 



node immediately above it, and those in the lower part of the 
intemode from the node below it. 

The organs of reproduction are very conspicuous by their colour 
and form. They are always situated on the leaves, the plants 
being very frequently monoecious. The antheridia (Fig. 61 B, a) 
are modified leaflets or the terminal cell of a leaf; they are spheri- 
cal and become red when mature. Their wall consists of 8 
" shields," i.e. of plate-like cells, 4 of which cover the upper half, 
and are triangular ; the 4 round the lower half, to which the stalk 
of the antheridia is attached, being quadrilateral, with sides of 
unequal length. The shields (Fig. 61 C) have dentated edges, 

FIG. 62. Chara fragilis : s apical cell ; FIG. 63. Oogonium of Chara : fc 

n, n nodal cells; in internodal cells; II, "crown"; u receptive spot; s sperma- 

II leaves ; r, r the cortical cells. tozoids. 

with the teeth fitting into one another, and their faces ornamented 
with ridges. From the centre of the internal face of each shield 
(C) a cylindrical cell, the manubrium, projects nearly as far as the 
centre of the antheridium ; at the inner end of each of the manu- 
bria a spherical cell, the capitulunt, is situated. Each capitulum 
bears six secondary capitula, from each of which four long coiled 
filaments (C, D) project into the cavity of the antheridium. 
These filaments are divided by transverse walls into from 100-200 
discoid cells, in each of which a biciliated, coiled spermatozoid is 
developed (D, E) from the nucleus. The spermatozoids escape 
from their mother-cell and are set free by the shields separating 
from one other. 



The female organ of reproduction (Fig. 61 -Z?, 63) is a small 
modified shoot, whose apical cell functions as an oogonium, its 
protoplasm forming the oosphere, which has a colourless receptive- 
spot at the summit (Fig. 63 ). The oogonium is situated on a 
nodal cell, from which 5 cells grow out in a circle and coil round 
the oogonium, covering it with a close investment. These cells 
divide once or twice at the top, so that 5 or 10 small cells are 
cut off, which project above the oogo- 
nium and form the so-called " crown " 
(Fig. 63 fc). The crown either drops off 
at fertilisation, or its cells separate to 
form a central canal for the passage of 
the spermatozoids. The wall of the 
oosphere l above the receptive spot be- 
comes mucilaginous, and allows the 
spermatozoid to fuse with the oosphere. 
The oospore, on germination (Fig. 64 sp), 
becomes a small filamentous plant of 
limited growth (Fig. 64 /, d, q, pi) the 
proembryo and from this, as a lateral 
outgrowth, the sexual generation is pro- 

The order is divided into two sub- 
orders : 

A. NITELLEJ;. The crown consists of 
10 cells ; cortex absent : Nitella, Toly- 

B. CHARES. The crown consists of 5 
cells ; cortex present : Tolypellopsis, Lam- 
prothamnus, Lychnotliamnus, Char a. 

Chara crinita is parthenogenetic ; in 
large districts of Europe only female 
plants are found, yet oospheres are 
formed capable of germination. 

About 40 species of fossilized Chara, 
determined by their carpogonia, are 
known in the geological formations from 
the Trias up to the present day. 

FIG. 64. Chara fragilis. Ger- 
minating oospore (sp); i, d, g, pi, 
form together the proembryo 
rbizoids (w") are formed at d ; 
w' the so-called tap-root; at g 
are the first leaves of the sexual 
plant, which appears as a lateral 

1 Before fertilisation the cosphere divides and cuts off at the base one or 
more cells (polar bodies?), termed " wendungszellen." 


Class 8. Phaeophycese (Olive-Brown Seaweeds). 

The Phaeophycea? are Alga?, with chromatophores in which the 
chlorophyll is masked by a brown colour (phycophcein). The pro- 
duct of assimilation is a carbohydrate (fucosan), never true starch. 
In the highest forms (FucAicece}, the thallus presents differentiation 
into stem, leaf, and root-like structures. The asexual reproduction 
takes place by means of zoospores. The sexual reproduction is 
effected by the coalescence of motile gametes, or by oogamous 
fertilisation. The swarm-cells are monosymmetric, each moved 
by two cilia which are true protoplasmic structures, and generally 
attached laterally (Fig. 65). The PhaaophyceaB are almost entirely 
salt-water forms ; a few species of Lithoderma live in fresh water. 

The class is divided into two families : 

1. PH^OSPORE^: : 1 Sub-Family, Zoogonicse; 2 Sub- Family, 

2. CYCLOSPORE.S! : Fucaceaa. 

Family 1. Phseosporese. 

The family consists of multicellular plants, whose cells are 
firmly united together to form a thallus ; this, in the simplest 
cases, may be a branched filament of cells (Ectocarpus), or, in the 
highest, may resemble a stem with leaves (Laminariacece) , while 
all transitional forms may be found between these two. The 
thallus grows by intercalary divisions (e.g. Ectocarpiis\ or by an 
apical cell (e.rj. Sphacelaria) ; pseudo-parenchymatous tissue may 
sometimes be formed by cells, which were originally distinct, 
becoming united together. The size of the thallus varies ; in 

some species it is quite small almost 
microscopical, while in the largest it 
is many metres in length. 

The vegetative cells in the lower 
forms are nearly uniform, but in those 
which are moi'e highly developed 
(Laminariacece and Fucacecc}, they are 
FIG. t>. bwaruispore of Cutler ia sometimes so highly differentiated that 

mechanical, assimilating, storing and 

conducting systems may be found ; the last named systems are 
formed of long cells with perforated, transverse walls, which bear 
a strong resemblance to the sieve-tubes in the higher plants. 
The colouring matter in the living cells (" phasophyl ") contains 


chlorophyll; but tliis is concealed by a brown (" phycophoein "), 
and a yellow (" phycoxanthin ") colouring material, and hence all 
these Algae are a lighter or darker yellow-brown. Starch is not 
formed. Asexual reproduction takes place, (1) by zoospores 
which arise in unilocular zoosporangia, and are raonosymmetric, 
with two cilia attached laterally at the base of the colourless 
anterior end (Fig. 65), the longer one being directed forwards 
and the shorter backwards ; or (2) by aplanospores (?). 

Sexnal reproduction has only been discovered in a few cases, and 
takes place by means of gametes (oogamous fertilisation perhaps 

FIG. 66. Ectocarpus tiliaulosug. I a-f 
A female gamete in the various stages 
of coming to rest. II A motionless 
female gamete surrounded by male 
gametes. Ill a^e Stages in the coal- 
escence of male and female gametes. 

PIG. 67. Zttiiardim'a collavis. A Male 
gametangia (the smaller celled) and female 
gatnetaiigia (the larger celled). C Female 
gamete. D Male gamete. JJ, E Fertilisation. 
F Zygote. G Germinating zygote. 

occurs in the Tilopteridse). The gametes have the same structure 
as the zoospores, and arise in multilocular gametangia ; these, like 
the zoosporangia, are outgrowths from the external surface, or 
arise as modifications from it. The conjugating gametes may be 
similar {e.g. Ectocarpus pusillus), or there may be a more or less 
pronounced difference of sex, an indication of which is found in 
Ectocarpus siliculosus (Fig. 60). When the gametes in this 
species have swarmed for a time, some, which are generally larger, 


are seen to attach themselves by one of the cilia, which by degrees 
is shortened to form a kind of stalk (compare the upper gamete in 
Fig. 66 II) ; these are the female gametes, which now become sur- 
rounded by a number of males endeavouring to conjugate with 
them, but only one succeeds in effecting fertilisation. The 
protoplasm of the two gametes coalesces (Fig. 66 III), and a 
zygote (e) is formed. The male gametes which do not conjugate 
may germinate, but the plants derived from them are much weaker 
than those produced by the zygotes. Strongly pronounced sexual 
differences are found in the Cutleriacese, in which order the male 
and female gametes arise in separate gametangia (Fig. 67 J.). The 
male gametes (Fig. 67 P) are much smaller than the female 
gamete (Fig. 67 C 1 ) ; the latter, after swarming for a short time, 
withdraws the cilia, and is then ready to become fertilised (Fig. 
67 I), E\ thus we have here a distinct transition to the oogamous 
fertilisation which is found in the Fucacere. Alternation of gene- 
rations is rarely found. 

1. Sub-Family. Zoogonicae. 
Reproduction by means of gametes and zoospores. 
Order 1. Ectocarpaceee. The thallns consists of single or 
branched filaments with intercalary growth, extending vertically 
from a horizontal, branched filament or a disc, but sometimes it is 
reduced to this basal portion only. Zoosporangia and gametangia 
(for fertilisation see Fig. 66) are either outgrowths or arise by 
the transformation of one or several of the ordinary cells. The 
most common genera are : Ectocarpus and Pylaiella. 

Order 2. Choristocarpacese. ChoristocarpuSjDiscosporaiigmm. 
Order 3. Sphacelariaceae. The thallus consists of small, 

parenchymatous, more or less ramified 
shoots, presenting a feather-like appear- 
ance. In the shoots, which grow by 
means of an apical cell (Fig. 68 S), a 
cortical layer, surrounding a row of 
central cells, is present. Sporangia and 
gametangia are outgrowths from the 
main stem or its branches. Sphacelaria, 
Chsetopteris are common forms. 

Order 4. Encoeliaceae. Punctaria, Aspero~ 

v , coccus, Plnillitis fascia, 

it IG. 6!-. Apex of tbe thallus 

of Clurtopteris plumosa. S Api- Order 5 - Striariaceae. Striana, Phlteo- 

cal cell. spora. 


Order 6. Dictyosiphonaceae. Dictyosiphon. 

Order 7. Desmarestiaceae. Desmarestia aculeata is common. 

Order 8. Myriotrichiaceae. Mijriotrichia. 

Order 9. Elachistaceae. Elachista fucicolu is a common epiphj'te on 
species of Fiicus. 

Order 10. Chordariaceae. The shoot-systems are often surrounded by 
mucilage. Chordaria ; Leathesia difformis occurs as rounded, brown-green 
masses of the size of a nut, generally attached to other Seaweeds. 

Order 11. Stilophoraceae. Stilophora rhizodes is common. 

Order 12. Spermatochnaceae. Spermatochnus paradoxus is common. 

Order 13. Sporochnaceae. Sporoclmus. 

Order 1-4. Ralfsiaceae. Ralfsia verrucosa is common as a red-brown incrus- 
tation on stoues and rocks at the water's edge. 

Order 15. Lithodermataceae. Some species of the genus Lithodcrma occur 
in fresh water. 

Order 16. Laminariaceae. The thallus is more or less leathery, 
and has generally a root-like lower part (Fig. 69) which serves to 
attach it, and a stalk or stem-like part, terminated by a large leaf- 
like expansion. Meristematic cells 
are situated at the base of the leaf, 
and from these the new leaves are 
derived. The older leaf thus 
pushed away by the intercalary 
formation of the younger ones, 
soon withers (Fig. 69). Gametes 
are wanting. Zoosporangia are 
developed from the lower part of 
a simple, few-celled sporangio- 
phore, which is an outgrowth from 
a surface-cell and has a large 
club-formed apical cell. The spo- 
rangia are aggregated into closely 
packed sori, which cover the lower 
part of the terminal leaf, or occur 
on special, smaller, lateral, fertile 
fronds (Alaria). Most of the 
species belonging to this order live 
in seas of moderate or cold tem- 
perature and occur in the most 
northern regions that have yet 
been explored, forming their organs 
of reproduction during the cold and darkness of the arctic night. 
Laminaria is destitute of a midrib and has only one terminal leaf. 

FIG. 69. Lammdria digitnta (much 
reduced in size) . 


L. digitata has a broad leaf, which, by the violence of the waves, is 
torn into a number of palmate strips (Fig. 69). L. saccharina has 
a small, undivided leaf. At aria has a midrib and special fertile 
fronds. A. esculenta occurs plentifully on the west coast of Nor- 
way and on the shores of Great Britain. Chorda filum, a common 
seaweed, is thick, unbranched, and attains a length of several 
metres, without any sti-ong demarcation between stalk and leaf. 
Some attain quite a gigantic size, e.g. Macrocystis pyrifera, whose 
thallus is said sometimes to be more than 300 metres in length. 
The Lessouia-STpecies, like the above, form submarine forests of 
seaweed on the south and south-west coasts of South America, 
the Cape, and other localities in the Southern Hemisphere. 

USES. The large Laminarias, where they occur in great numbers, are, like the 
Fuci, used for various purposes, for example, in the production of iodine and 
soda, and as ail article of food (Laminaria saccharina, Alaria exculenta, etc). 
Laminaria mu-L-harina contains a large quantity of sugar (maniiit) and is in 
some districts u-;ed in the preparation of a kind ol syrup ; in surgical operations 
it is employed for the distension of apertures and passages, as for instance the 
ear-passage. It is by reason of the anatomical peculiarities and structure of the 
cell-walls, that they are employed for this purpose. The cell-walls are divided 
into two layers, an inner oue which has very little power of swelling, and an 
outer one, well developed and almost gelatinous the so-called "intercellular 
substance'' which shrivels up when dried, but can absorb water and swell to 
about five times its &i/e. The stalks of Laininuria clustoiii are oflicinal. 

Order 17. Cutleriacese. The thallus is formed by the union 
of the originally free, band-shaped shoots. The growth is inter- 
calary. Sexual reproduction by the conjugation of male and 
female gametes. An asexual generation of different appearance, 
which produces zoospores, arises from the germination of the 
zygote. Cutleria, Zaiiardinia. 

Sub-Family 2. Acinetae. 

Branched, simple cell-rows with intercalaiy growth. The 
organs of reproduction are partly uni- and partly multi-cellular ; 
in the unicellular ones a cell without cilia is formed, which may be 
destitute of a cell-wall, but has one nucleus (oosphere ?), or which 
has a cell-wall and contains several (generally four) nuclei 
(aplanospores ?) ; in the multicellular, monosymmetric swarm-cells 
with two cilia (spermatozoids r*) are formed. The fertilisation has 
not been observed. 

Order 1. Tilopteridaceae. Haplospora, Tilopteris. 


Family 2. Cyclosporeae. 

The individuals are rnulticellular, with growth by an apical 
cell. The thallus often bilateral is differentiated into a root- 
like structure (attachment-disc), and stem, sometimes also into 
leaves (Sargassum). Sometimes a differentiation occurs into 
various tissue-systems, viz. an external assimilating tissue, a storing 
tissue, a mechanical tissue of thickened, longitudinal, parenchyma- 
tous, strengthening cells, and a conducting tissue of sieve-cells, or of 
short sieve-tubes with perforated walls. Colouring material, as 
in Phaeosporege. Vegetative reproduction can only take place by 
means of detached portions of the thallus (Sarg>issum\ which are 
kept floating by means of bladders (Fig. 70 A, a, Fig. 72). Zoo- 
spores are wanting. 

The sexual reproduction takes place by oogamous fertilisation. 
The oogonia and antheridia are formed inside special organs 
(conceptacles), and are surrounded by paraphyses. The concep- 
tacles (Fig. 70 JJ, Fig. 71 b) are small, pear-shaped or spherical 
depressions, produced by a special ingrowth of the surface cells 
of the thallus, and their mouths (ostioles) project like small warts ; 
they are either situated near the end of the ordinary branches of 
the thallus (Fucus serratus, Fig. 71 a) which may be swollen on 
this account (Fucus vesiculosus, Fig. 70 A, 6), or on special short 
branches (AscopTvyllum, Sargassum). The vertical section of a 
coiiceptacle is seen in Fig. 70 B (see also Fig. 71 &) where, in 
addition to the paraphyses, oogonia only are seen (F. vesiculosus is 
dioecious male plant, yellow-brown; female plant, olive-brown); 
but in some species antheridia, together with oogonia, are pro- 
duced in the same coiiceptacle. The oogonia are large, almost 
spherical cells, situated on a short stalk, in each of which are 
formed from 1-8 (in Fucus, 8; in Ascophyllum, 4; in Halidrys, 1 ; 
in Pelvetia, 2) rounded, immotile oospheres. The wall of the 
oogonium ruptures, and the oospheres, still enclosed in the inner 
membrane, are ejected through the mouth of the conceptacle, and 
float about in the water, being finally set free by the bursting of the 
inner membrane. The antheridia are oblong cells (Fig. 70 G, a), 
many of which are produced on the same branched antheridio- 
phore (Fig. 70 C) ; the numerous spermatozoids are provided 
with 2 cilia and are very small (Fig. 70 D, two antheridia sur- 
rounded by spermatozoids, one being open). The spermatozoids, 
still enclosed by the inner membrane of the antheridium, are 



similarly set free, and fertilisation takes place in the water, 
numerous spermatozoids collecting round the oosphere (Fig. 70 E), 
which is many times larger, and by their own motion causing it' to 
rotate. After fertilisation, the oospore surrounds itself with a 

& :^Mg^*fr*s! 




FIG. 70. Fttcits vesiculosus. A Portion of thallus with swimrciDg bladders (n) and 
conceptacles (6). S Section of a female conceptacle ; li the mouth; 31 the inner cavity; 
s ooponia. C Antheridiophore; a antheridium; y sterile cells. D Anthcridia out of which 
the eparmatozoicls are escaping. E Fertilisation. F Germinating oospore. 

cell-wall and germinates immediately, attaching itself (Fig. 70 
to some object, and by cell-division grows into a new plant. 



Order 1. Fucaceae. The following species are common on our 
coasts : Fucus vesiculosus (Fig. 70) has a thallus with an entire 
margin, and with bladders arranged in pairs ; F. serratus (Fig. 71) 
without bladders, but with serrated margin ; Ascophijllum nodosiim 
has strap-like shoots, which here and there are swollen to form 
bladders ; Halidrys siliquosus has its swimming bladders divided 
by transverse walls; Himanthalia Zorea, which is found on the west 
coast of Norway, and the south, coast of England, has a small 
perennial, button-shaped part, from the centre of which proceeds 
the long and sparsely branched, strap-like, annual shoot, which 

FIG. 72. Sargafsum baccifernm. A 
portion of the thallus, natural size. 

FIG. 71. Fucus serratus. a Portion of a male plant -which has been exposed to the 
action of the open air for some time ; small orange-yellow masses, formed by the anther- 
idia.are seen outside the mouths of the male cnneeptacles (nat.size). b Cross section through 
the end of a branch of a female plant, showing the female conceptacles ( x 4). 

bears the conceptacles. The Gulf-weed (Sargassiim bacciferum, 
Fig. 72) is well known historically from the voyage of Columbus ; 
it is met with in large, floating, detached masses in all oceans, 
and is found most abundantly in the Atlantic, off the Canary 
Islands and the Azores, and towards the Bermudas. The 
stalked, spherical air-bladders are the characteristic feature of 
this genus. The thallus is more highly developed than in Fucus, 
and there is a contrast between the stem and leaf-like parts. The 


portions which are found floating are always barren, only those 
attached are fertile. 

USES. The Fucaeeae, like the Laminariaceae, are used as manure (the best 
kinds being Fucus vtsicnlosu* and Aicophijllum nodosum), for burning to pro- 
duce kelp, and as food for domestic animals (^scopliyllum nodosum is especially 
used for this purpose). 

Class 9. Dictyotales. 

The plants in this class are multicellular, and brown, with 
apical growth, new cells being derived either from a flat apical 
cell, or from a border of apical cells. The thallus is flat, leaf- or 
strap-shaped, attached by haptera, which are either found only 
at the base, or on the whole of the lower expansion of the thallns. 
The cells are differentiated into the following systems of tissues : 
an external, small-celled layer of assimilating cells, generally one 
cell in thickness, and an internal, large-celled layer of one or only 
a few cells in thickness, forming the mechanical and conducting 
tissues. All the reproductive cells are motionless. Asexual re- 
production by naked, motionless spores (tetraspores) which are 
formed 1-4 in each tetrasporangium, the latter being outgrowths 
from the surface cells of special, sexless individuals. Zoospores 
are wanting. The sexual organs are of two kinds, oogonia and 
antheridia, which are formed from the surface cells, either on the 
same or different individuals. The oogonia are spherical or oval, 
and are generally placed close together ; each contains one 
oosphere, which on maturity is ejected into the surrounding 
water, and is then naked and motionless. The antheridia are 
formed of longitudinal cells, united in groups, whose contents 
by repeated divisions transverse and longitudinal are divided 
into a large number of small, colourless, motionless sperm at ia - 
round or elongated which are set free by the dissolution of the 
wall of the antheridium. The process of fertilisation has not 
yet been observed. 

The Dictyotales, in having tetraspores and spermatia, deviate 
considerably from the Pha9ophycea3, but may be classed near to 
the Tilopterida?, in which there are asexual spores with 4 cell- 
nuclei, which may be considered as an indication of the formation 
of tetraspores. 

Order 1. Dictyotaceae. Dictyota dichotoma which has a thin, regularly 
dichotoruously divided thallus, occurs on the coasts of the British Isles 
Padina is found on the south coast. 

BANG10IDE.E. 77 

Class 10. Rhodophyceae (Red Seaweeds). 

The plants comprised in this class are multicellular ; they are 
simple or branched filaments, or expansions consisting of 1 to 
several layers of cells; the thallus maybe differentiated (as in 
many Floridece), to resemble stem, root, and leaf. The cells con- 
tain a distinctly differentiated nucleus (sometimes several), and 
distinct chromatophores, coloured by rhodophyll. The chloro- 
phyll of the chromatophores is generally masked by a red colour- 
ing matter (phycoerythrin), which may be extracted in cold, fresh 
water ; or rarely by phycocyan. Pyrenoids occur in some. Starch 
is never formed in the chromatophores themselves, but a modifi- 
cation Floridea? starch may be found in the colourless proto- 
plasm. Asexual reproduction by motile or motionless spores 
(tetraspores) which are devoid of cilia and of cell-wall. Swarm- 
spores are never found. 

Sexual reproduction is wanting, or takes place by the coales- 
cence of a spermatium and a more or less developed female cell. 
The spermatia are naked masses of protoplasm, devoid of cilia and 
chromatophores. The female cell (carpogonium) is enclosed by a 
cell-wall, and after fertilisation forms a number of spores, either 
with or without cell-walls (carpospores), which grow into new 

The Rhodophycea3 may be divided into two families : 

Family 1. Bangioidea.. 

The thallus consists of a branched or unbranched cell-filament, 
formed of a single row or of many rows of cells, or of an expan- 
sion, one or two layers of cells in thickness, but without conspic- 
uous pores for the intercommunication of the cells. The growth of 
the thallus is chiefly intercalary. The star-like chromatophores 
contain chlorophyll and are coloured blue-green with phycocyan, 
or reddish with phycoerythrin ; all these colouring' matters are 
occasionally found in the same cell (Bangia-species). Asexual 
reproduction by tetraspores, without cilia, but capable of amoeboid 

Sexual reproduction is wanting, or takes place by the coalescence 
of a spermatium with a carpogonium, which is only slightly differ- 
entiated from the vegetative cells, and is devoid of a trichogyne. 



The carpospores are destitute of cell-wall and arise directly by 
the division of the fertilised oosphere. The Bangioidete occur 
chiefly in salt water. 

Order 1. Goniotrichacese. The thallus consists of a branched cell- 
filament without rhizoids. Tetraspores are formed directly from the entire 
contents of the mother-cell, without any preceding division. Fertilisation 
unknown. Asterocystis, Goniotrichum. 

The GoniotrichacecK, through the blue-green Asterocystis, are allied to the 
Myxophycese, and through Goniotrichum to the Porphyraeeee. 

Order 2. Porphyraceae. The thallus is formed of an expansion consisting 
of a layer of 1-2 cells, which, at the base, are attached to the substratum by 
means of a special form of haptera (Porphyra, Diploderma) ; or of uubranched 
(very rarely slightly branched) filaments, attached at the base by haptera 
(Bangia) : or it extends from a prostrate cell-disc (various species of Erythro- 
trichia). Tetraspores are formed after one or more divisions of the mother-cell, 
either from the whole or only a part of its contents ; they possess arnceboid 
movements, or have a jerky, sliding-forward motion. The antheridia have 
the same appearance as the vegetative cells, but divide several times, and 
several spermatia are formed, either simultaneously from the whole contents 
(Porphyra, Bangia), or the spermatia are successively formed from a part of 
the contents of the antheridium (Erythrotrichia). The carpogonium is with- 
out a trichogyne, but the oosphere has a colourless spot which may sonae- 
times rise a little above the surface of the thallus, and may be considered 
as an early stage in the development of the trichogyne. The spermatia form 
a canal through the membrane of the carpogonium, and their contents coal- 
esce with the oosphere at its colourless spot. The fertilised oosphere divides 
on germination into a number of carpospores, which are set free as naked, 
motionless masses of protoplasm, which grow and give rise to new individuals 
(alternation of generations). 

Family 2. Floridese. 

The thallus has one or more apical cells, grows principally by 
apical growth, and may be differentiated into root, stem, and leaf. 
The chromatophores vary in form, but have a red or brownish 
colour, due to chlorophyll and phycoerythrin. Asexual repro- 
duction by motionless tetraspores, which generally arise by the 
division into four of the contents of the tetrasporangium. The 
carpogonium has a trichogyne, and the carpospores, which are 
formed indirectly from the fertilised oosphere, possess a cell- 

The thallus may assume very different forms. In the simplest 
species it is filamentous and formed of single, branched rows of 
cells (Callithamnion, etc., Fig. 73). Ceramium has a filamentous, 
thallus, generally dichotomously forked (Fig. 75), or sometimes 


pinnately branched, which, at the nodes, or throughout its entire 
length, is covered by a layer of small cortical cells. Polysiphonia 
(Fig. 74) has a filamentous, much branched thallus, made up of 
a central cylindrical cell, surrounded by a layer of other cells, 
cortical cells, which in length and position correspond to the 
central ones. In many of the Red Alga? the vegetative organs 
are differentiated into stems and leaves, the former having, as in 
Chara, unlimited growth in length, whilst the latter soon attain 
their full development. Choiidrus has a fleshy, gelatinous thallus, 
without nodes ; it is repeatedly forked into flat branches of vary- 
ing thickness. Farcellaria has a forked thallus with thick branches 

FIG. 73. CaUitJmmnion elegans : a a plant with 
tetraspores ( x 20) ; b apex of a branch with 
tetraspores ( x 250). 

Fis. 74. Polj/si'pTioni'a variegata : 
a a portion of a male plant with 
aiitheridia; b spermatia ; c trans- 
verse section of thallus. 

and without nodes. The thallus of Delesseria (Fig. 76) consists of 
branches, often bearing leaf-like structures, with a midrib and 
lateral ribs springing from it. These ribs persist through the 
winter, and at the commencement of the succeeding period of 
vegetation the lateral ribs become the starting points for new- 
leaves. In Corallina the thallus is pinnately branched, and 
divided into nodes and internodes. The name has been given to 
this genus from the fact that the thallus is incrusted with car r 
bonate of lime to such a degree that it becomes, very hard, and the 



whole plant adopts a coral-like appearance. Other genera which 
are similarly incrusted, and have a leaf-like or even crustaceous 
thallus (such as Melol>esia, Lithothamnion), are included in this 

In some instances the cells of the thallus may be found differ- 
entiated into more or less well defined tissues, so that it is possible 
to find special assimilating, mechanical, and conducting tissues, 
the last named in some cases having the double function of con- 
ducting and of serving as a reservoir in which starch is found as a 
reserve material. The cells of the Floridese, which are formed by 
the division of a mother-cell into two daughter-cells of unequal 

FIG. 75. Ccramium diaplianum (uat. size). 

FIG. 76. Delesseria sanguinea (about 

size, have always larger or smaller pits in the cell-walls, and the 
thin cell- wall separating two pits from each other is perforated by 
a number of small holes. These pits are particularly developed in 
the conducting tissues, but sieve-tubes are very rarely to be found. 

Tetraspores may be wanting (e.g. Lemanea) or may often arise 
on special, non-sexual individuals. In some (e.g. Bntrachospir- 
mum) only one tetraspore is formed in each tetrasporangium, but 
the number is generally four, which may be formed tetrahedrally 
(Fig. 73) or by divisional walls perpendicular to each other, or 
even in a single row. The tetrasporangia in some species are free 
(Fig. 73), but in the majority they are embedded in the thallns. 

The sexual reproduction (discovered by Thuret and Bornet, 



1867) differs in the essential points from that of all other 
plants, and approaches most nearly to the sexual reproduction 
of the Bang-widen;. The sexual cells are developed from the 
terminal cells (never nodal cells) of the branched cell-filaments, 
which constitute the thallus. The mother-cells of the spermatia 
(spermatangia) are generally arranged in a group, in the so-called 
antlieri'Hci (Figs. 74, 77 A, a). On becoming ripe the membrane 
of the spermatangium ruptures and the spermatia emerge as 
spherical or ovoid, naked (a little later they may possess a cell- 
wall) masses of protoplasm which are not endowed with the power 
of motion, and hence are carried passively by the current of the 

FIG. 77. A Lejolisia mediterranea : rhaptera; s longitudinal section through a cystocarp ; 
p the empty space left by the liberated spore (0- -B-.E Nemalion multifidum: a antheridia ; 
b procaruium with trichogyne, to which two spermutia are adhering. 

water in which they may happen to be, to the female cell. This 
latter is analogous with the oogonium of the Green Algre. The 
female reproductive organ is termed the procarpium, and consists 
of two parts, a lower swollen portion the carpogonium (Fig. 77 b 
in A and B) Avhich contains the cell-nucleus, and an upper 
filamentous prolongation the trichogyne (Fig. 77 B) which is 
homologous with the colourless receptive spot of the oosphere of 
the Green Algre, and the Porphyracece. In the sexual reproduc- 
tion of the majority of the Floridea?, a very important part is 
played by certain special cells, rich in cell-contents the auxiliary 
u. B. G 


cells. These are either dispersed in the interior of the thallns, 
or are arranged together in pairs with the cell-filament which 
bears the carpogonium, and are generally united with this to 
form an independent multicellular procarpium. The sperrnatia 
attach themselves firmly to the trichogyne and surround them- 
selves with a cell-wall. The dividing wall at the point of contact 
is perforated, and the nucleus of the spermatium probably travels 
through the trichogyne to the swollen part of the procarpium 
the carpogonium and fuses with its nucleus. After fertilisation 
the trichogyne withers (Fig. 77 0), but the lower portion'of the 
procarpium, constituting the fertilised oosphere, grows out and 
forms in various ways, first a tuft of spore-forming filaments 
known as gonimoblasts, and finally the carpospores. These latter 
form a new asexual generation (compare the germination of the 
oospore of OEdogonium and Goleochcete'). 

The gonimoblasts may arise in three ways : 

1. In the Nemalionales, branched filaments grow out from the oosphere and 
form an upright, compressed or expanded tuft of spore-forming filaments. 

2. In the Cryptoneiuiales, several branched or uubranched filaments (ooblas- 
tema-filaments) grow out from the oosphere, and conjugate in various ways with 
the auxiliary cells. The gonimoblasts are then formed from the single cells 
produced by the conjugation. 

3. In the Gigartinaies and Rhodymeniales the oosphere conjugates with an 
auxiliary cell by means of a short ooblasterna-filament, and from this auxiliary 
cell a gonimoblast is produced. 

The motionless carpospores, which sometimes in the early stages are naked, 
and afterwards invested with a cell-wall, are developed from the terminal cells 
(and perhaps also from some of the other cells) of the branches of the goniuio- 
blast. The gonimoblasts constitute sharply defined parts of the plant in which 
the carpospores arise. These parts are called cystocarps and are either naked 
(Fig. 77 E), or surrounded by a covering (pericarp or involucre, Fig. 77 A) 
formed in different ways. On this account the Florideac were formerly divided 
into GYMNOSPORE/E (Batrachospermum, Nemalion, Cerainiuni, etc.) and ANOIO- 
SPORE.E (Fnrcellaria, Lejolisia, Delesseria, Melobesia, etc.). 

The Floridese are divided into four sub-families : 

Sub-Family I. Nemalionales. The fertilised oosphere produces directly 
the gonimoblast. 

Order 1. Lemaneaceae. Algas of brownish colour and living in fresh 
water. They lack tetraspores, and the very sparingly bra7iched fertile filaments, 
composed of many rows of cells, grow out from a pro-embryo, which consists of 
a single row of cells bearing branches. Lemanea Jluviatilis, often found on 
rocks and stones in quickly flowing streams. 


Order 2. Helminthocladiacese. Tetraspores are generally wanting (e.g. in 
Nentalioit) or arise one in each tetrasporangiuui (e.g. Batrachospermum) and it 
is only in Liagora that four cruciate tetraspores are formed. CMntranxia 
corymbifera consists of simple, branched cell-rows, and is an independent species. 
Several other Cliantransia-forms, living in fresh water, are "proembryos" of 
species of the genus Batrachotpermum. The germinating carpospore grows out 
into filaments and forms a so-called proembryo which, if not shaded, attains 
only a small size, but when growing in shady situations presents a much greater 
development. These highly developed proembryos have been described as 
species of Chantransia. The proembryo can reproduce by division, or by tetra- 
spores which are developed singly in the sporangia; in B. vayum and B. spuru- 
lunx which do not possess fully developed female reproductive organs, the pro- 
embryos serve almost entirely to reproduce the species. The young Batracho- 
spermum-plant arises from the end of an upright filament of the proembryo. 
The proembryo is generally persistent, and continually produces new Batraeho- 
spennums. These latter bear the sexual reproductive organs and also whorls of 
branches : the central row of cells is enclosed by cells growing from the base of 
the whorls of branches, and from these cortical cells secondary proembryos are 
developed. In this alternation of shoots there is really no alternation of 
generations, since the proembryo and the shoots with the sexual reproductive 
organs are parts of the same thallus. 

Several species of Batrachospermum have a bluish green or verdigris colour. 
Nemalion multijidum has a brown-red thallus, slightly branched, which is 
attached to rocks near the water's edge. 

Order 3. Chaetangiaceae. Galuxaura Las a thallus thickly incrusted with 

Order 4. Gelidiaceae. Naccaria, Gelidium. 

Sub-Family 2. Gigartinales. The fertilised auxiliary cell grows towards 
the thallus, to produce the gonimoblasts. Procarpia generally present. 

Order 5. Acrotylaceae. Acrotylus. 

Order 6. Gigartinaceae. 6-igartina, Phyllophora, Ahiifeltia ; Chondrus 
crispus, with dark red, dichotomously branched thallus, is common on the 
coasts of Scandinavia and Great Britain. 

Order 7. Rhodophyllidaceae. Ehodophyllis, Euthora ; Cystoclonium pur- 
purascens is common, and sometimes the ends of its branches may be modified 
into tendril-like haptera. 

Sub-Family 3. Rhodymeniales. The fertilised auxiliary cell forms the 
gommoblast on the side away from the thallus. Procarpia are abundantly 

Order 8. Sphaerococcaceae. Gracilariu. 

Order 9. Rhodymeniaceae. Rhodytneiua palinata is a common species. 
Lomentaria, Chylocladla, Plucamium. 

Order 10. Delesseriaceas. Deletseria sanguined ; D. alata and I), sinuosa 
aie handsome forms which are not uncommon. 

Order 11. Bonnemaisoniaceae. Bonnemaisonia. 

Order 12. Rhodomelaceae. Rhodoinela, Odonthalia ; Polysiphonia, of 
which many species are to be found on the coasts of Great Britain, has a 
filamentous, richly branched thallus consisting of a central row of cells sur- 

84 FUNGI. 

rounded by a varying number of cortical cells of similar size the so-called 
" siphons." 

Order 13. Ceramiaceae. Pretty Algfe, often branched dichotomously, or 
unilaterally pinnate. Spermothamition, Grijfithsiu, Callithamnivn, Ccramium, 

Sub-Family 4. Cryptonemiales. The cells formed by the coalescence 
of the auxiliary cells and the ooblastema-filameuts, produce the gouimoblasts. 
The carpogunium-jilaments and the auxiliary cells are scattered siugly in the 

Order 14. Gloiosiphoniaceas. Gloiopeltis. 

Order 15. Grateloupiaceas. Halymenia, Cryptoncmia. 

Order 16. Dumontiaceas. Dumontiii, DuJretniaya. 

Order 17. Nemastomacese. Furcellaria, which has dichotomously 
branched, round shoots, is common on the coasts of Great Britain. 

Order 18. Rhizophyllidaceae. PolyiJes, Rhizophyllis. 

Order 19. Squamariaceas. The Alga- belonging to this order form crust- 
like coverings on stones, mussel-shells, and on other Algffi, but are not them- 
selves incrustated : Petrocflis, Cruoria, Peyssonellia, 

Order 20. Corallinaceas. Partly crustaceous, partly erect, branched Alga?, 
thickly iucrusted with lime, so that a few species (LitliotJiamnia, also called 
NulUpora') occur in fossilized condition from Jurassic to Tertiary periods. 
3Jelubaia, Lithophyllum, Lithothamnion, Corallina. 

USES. " Carragen " is the tliallus of Cfiondms cripus (Irish 
Moss) and Gigartina mamillosa. It is a common article of food on 
the coasts of Ireland, and swells to a jelly when cooked. It is 
officinal. Rhodymenia palmetto, is generally eaten as food in 
Ireland and in some places on the west coast of Norway ; it is 
also used as food for sheep and hence is termed " Sheep-seaweed. " 
Agar-Agar is the jelly obtained from species of Gelid ium and 
Gigartina growing in China and Japan. 

Sub-Division III. FUNGI. 

Mode of Life. The Fungi have no chlorophyll, and are thus 
unable in any stage of their existence to assimilate carbon ; they 
must therefore live as saprophytes or parasites. There is, however, 
no strong line of demarcation between these ; many Fungi com- 
mence as true parasites, bat only attain their full develop- 
ment upon or in dead plants or animals (Rhytisma, Empusa). 
Many saprophytes may occasionally appear as parasites, and are 
then designated "facultative parasites ' (Nectria cinnabarinn, 
Lophodermium piiumtri), in contradistinction to those AY Inch only 

FCNGI. 85 

appear as parasites, " obligate parasites" (Mildew, Brand- and Rust- 
Fungi, Cordyceps). 

The parasites which live on the surface of the host-plant are 
termed epiphytic (Mildew, Fiisicladium} ; and those living in its 
tissues are termed endopliytic (Ustilago, Peronospora). Epizoio 
(Oidium tonsnrans, Laboulbenia) and endnzoic Fungi (Cordyce/'>s i 
Eiitomnplithora), are distinguished, in the same manner, as those 
which live on the surface or in the interior of animals. The Funo-i 


designated pathogenic are especially those which produce disease 
in human beings and in animals. 

Most of the diseases of plants are attributed to the parasitic 
Fungi. These force their way into the host-plant by piercing the 
outer wall of the epidermis, as in the Potato-disease; or by grow- 
ing in through the stomata, e.g. the summer generations of the 
Rust of Wheat ; or they can only penetrate through a wound, 
e.g. Nectria. Some effect an entrance into the host- plant by the 
secretion of a poisonous matter or ferment, which, softens and 
destroys the cell-walls (Sderotinia). Some Yeast and Mould 
Fungi secrete ferments (enzymes), which, for example, convert 
cane-sugar into a sugar capable of fermentation. 

The relation of the parasitic Fungus to the host-plant is mainly 
of two kinds. In the one case, the cell-contents are desti'oyed, 
the protoplasm is killed, and the cellular tissue becomes discoloured 
and dies (Peronospora, Armillaria mellea, Polyporus) ; in the 
other case, the parasite has an irritating effect on the cellular 
tissue, whereby the affected organ grows more rapidly and be- 
comes larger than normal, producing hypertrophy. Such malfor- 
mations are termed Fungi-galls (Mycocecidia) ; in this manner 
" witches' brooms " are produced by jEcidium, " pocket-plum " by 
Taphrina, and other deformities by Exobasidium and Cystnpus 
candi'lus. This hypertrophy may either be produced by a vigorous 
cell-multiplication, which is most frequently the case, or by the 
enlargement of the individual cells (Synchytrium, Calyptospora). 
The relation between host and Fungus among the Lichens is of 
a very peculiar nature, termed " symbiosis." 

Vegetative Organs. The vegetative parts of a Fungus are 
termed its mycelium. 1 This is formed of a mass of long, cylindrical, 
branched cells resembling threads (and hence termed hyphen), 
which have a continued apical growth. The mycelium, in its 
early development, shows a well-marked difference between the 
1 From the Greek /j.vKt)s = Fungus, hence " myc 'logy." 

86 FUNGI. 

two main groups of true Fungi : in the Pliy corny cetes, or Algal 
Fungi, the mycelium has no transverse walls, and is therefore 
unicellular, while in the Mesomycetes and My corny cetes it is pro- 
A'ided with dividing walls, which grad^^ally arise during growth, in 
the youngest hypha? ; intercalary transverse walls may also be 
formed at a later period. In the hyphas of some of the Higher 
Fungi (Hymenomycetes), connections may be formed between two 
contiguous cells of the same hypha, by a protuberance growing out 
from an upper cell just above the transverse wall, and forming a 
junction with the cell below. These are known as clamp-connec- 
tions; they appear to be of use in affording communication be- 
tween the two cells. 

The hyphse of Fungi, where they come in contact with one 
another, often grow T together, so that H-formed combinations (fu- 
sions) are produced, which give rise to very compact felted tissue. 
When the hypha? are not only closely interw r oven, but also united 
and provided with many transverse walls, the mycelium assumes 
the appearance of a tissue with isodiametric cells, and is then 
termed pseudo-parencJiyma. The hyphse-walls are sometimes very 
much thickened, and composed of several layers, and the external 
layers, by the absorption of water, may often swell very much 
and become mucilaginous. In some instances the walls are colour- 
less, in others coloured, the most frequent colour being brown. 
The cell-contents may also be coloured, and in that case are gene- 
rally yellow; this colour is chiefly connected with the fat (oil) 
which may be found in abundance in the Fungi, whilst starch is 
invariably absent in all the true Fungi. 

The mycelium assumes many different forms ; sometimes it 
appears as a thread-like, cobwebby, loose tissue, less frequently as 
firm strands, thin or thick membranes, horn-like plates or tuber- 
like bodies. The thread-like mycelium may, in the parasitic Fungi, 
be intercellular or intracellular, according as it only extends into 
the interstices between the cells or enters into the cells proper. 
In the first case there are generally found haustoria, or organs of 
suction (e.g. among the Peronosporacece ; Taphrina, on the contrary, 
has no haustoria) ; but haustoria are also found among the 
epiphytic Fungi (e.g. Erysiphacea^). Intracellular naycelia are 
found in the Rust-Fungi, in Claviceps purpurea, EutomopJithora, etc. 
In spite of its delicate structure, this mycelium may live a long 
time, owing to the circumstance that it continues to grow peri- 
pherally, while the older parts gradually die off ("fairy rings"). 

FUNGI. 87 

String-like mycelia may be found, for example, in Phallus, 
Coprinus, and are formed of hyphse, which run more or less 
parallel to each other. Membrane-like mycelia are chiefly to be 
found in Fungi growing on tree-stems (Polyporacea? and Agari- 
cacea3) ; they may have a thickness varying from that of the 
finest tissue-paper to that of thick leather, and may extend for 
several feet. The peculiar horny or leather-like strands and plates 
which, for instance, appear in Armillaria mellea, are known as 
Rhizomorpha ; they may attain a length of more than fifty feet. 
The tuber-like mycelia or sclerotia play the part of resting mycelia, 
since a store of nourishment is accumulated in them, and after a 
period of rest they develope organs of reproduction. The sclerotia 
are hard, spherical, or irregular bodies, from the size of a cabbage 
seed to that of a hand, internally white or greyish, with a brown 
or black, pseudo-parenchymatous, external layer. Sclerotia only 
occur in the higher Fungi, and are found both in saprophytes, e.g. 
Coprinus, and in parasites, e.g. CJaviceps (Ergot), Sclerotinia. 

Reproduction. SEXUAL REPRODUCTION is found only among 
the lower Fungi which stand near to the Algae, the Algal-Fungi, 
and takes place by the same two methods as in the Algte, namely 
by conjugation and by the fertilisation of the egg-cell in the 

The majority of Fungi have only ASEXUAL reproduction. The 
most important methods of this kind of reproduction are the 
sporangia- fructification and the conidio-fructijication. 

In the SPORANGIO-FKUCTIFICATION the spores (endospores) arise 
inside a mother-cell, the sporangium (Fig. 80). Spores without a 
cell-wall, which move in water by means of cilia and hence are 
known as swarmspores or zoospores, are found among the Oomycetes, 
the sporangia in which these are produced being called swarm- 
sporangia or zoosporangia (Figs. 86, 87, 91, 94). 

In the CONIDIO-FRUCTIFICATION the conidia (exospores) arise on 
special hyphse (conidiophores), or directly from the mycelium. 
When conidiophores are present, the conidia are developed upon 
them terminally or laterally, either in a basipetal succession 
(in many Fungi, for example in Penicillium, Fig. Ill, Erysiphe, 
Cystopus), or acropetally (in which method the chains of conidia 
are often branched ; examples, Pleospora vulgaris, Hormodendron 
cladosporioides) . All conidia are at first unicellular, sometimes at 
a later stage they become two-celled or multicellular through the 
formation of partition-walls (Piptocephalis) . The conidia with 

88 FUNGI. 

thick, brown cell-walls, and contents rich in fats (resting conidia), 
can withstand unfavourable external conditions for a much longer 
period than conidia with thin walls and poor in contents. 

The SPORANGIA arise either from the ordinary cells of the 
mycelium (Protomyces\ or are borne on special hyphre. They are 
generally spherical (Mucor, Fig. 80 ; Saprolegniacese), egg-, pear-, 
or club-shaped (Ascoriiycetes), more rarely they are cylindrical or 
spindle-shaped. While among the Phycornycetes the size, form, 
and number of spores are indefinite in each species, in the Ascomy- 
cetes the sporangia (asci) have a definite size, form, and number 
of spores. The spores of the Ascomycetes are known as asco- 

The sporangio-fructification is found under three main forms. 

1. FREE SPORANGIOPHORES which are either single (Mucor, Fig. 
78), or branched (Thamnidium). 

2. SPORANGIAL-LAYERS. These are produced by a number of 
sessile or shortly-stalked sporangia, being formed close together 
like a palisade (Taphrina, Fig. 105). 

3. SPORANGIOCARPS. These consist usually of many sporangia 
enclosed in a covering, they are found only in the Carpoasci, and 
are also known as ascocarps. The parts of an ascocarp are the 
covering (peridiutn), and the hymenium, which is in contact with 
the inner wall of the peridium, and is generally made up of asci, 
and sterile, slender hyphje. The latter either penetrate between 
the asci and are branched and multicellular (paraphyses, Figs. 
103 d, 123, 125, 129), or clothe those parts of the inner wall which 
bear no asci (Periphyses ; among many peronocarpic Ascomycetes, 
e.g. Chwfomium, Sordaria, Stictosphce.ra lioffmanni). The ascocarps 
are produced directly from the mycelium, or from a stroma, that 
is a vegetative body of various forms, in which they may be 
embedded (Figs. 116 B, C). 

Among the conidio-fructifications there are, in the same way, 
three divisions. 

1. FREE CONIDIOPHORES (Fig. 109). The form of the conidiophores, 
the shape, and number of its spores are various. Iii the most 
highly developed Fungi, the Basidiomycetes, there are, however, 
special more highly developed conidiophores, the basidia, which 
have a definite form ana spores of a definite shape and number. 
The conidia borne on basidia are called busidiozpores. 

2. CONIDIAL-LAYERS. (a) The SIMPLEST case of this is found 
when the conidiophores arise directly from the mycelium, parallel 

.FUNG[. 89 

to one another, and form a flat body (e.g. Exobasidium vaccinii, 
Hypochnus ; among the Phycomycetes, Empusa nniscce and Cystopus). 
(b) In a HIGHER form the conidial-layers are thick, felted threads 
(stroma) inserted between the mycelium and the hymenium (i.e. 
the region of the conidiophores). Examples are found in a section 
of the Pyrenomycetes (Fig. 122). (c) The HIGHEST form has the 
basiJiaJ -layer, that is a conidial-layer with more highly developed 
conidiophores (basidia). The basidial-layer, with stroma, and 
the hymeniurn (region of the basidia), forms the basidio-fructifica- 
tion, which is branched in the Clavariacea?, and hat-shaped in 
other Hymenomycetes (in these groups the hymenium is confined 
to the lower side of the pileus). 

The hymenium of the conidial-layer and basidial-layer is com- 
posed entirely of conidiophores, or of conidiophores and sterile 
hyphre (paraphyses) which are probably always unicellular. 
Paraphyses are found in Entomophthora radicans, and in certain 
Basidiomycetes (e.g. Corticium'). 

3. CONIDIOCARPS (pycmdia). A special covering surrounds the 
conidia-foi-ming elements. The inner side of this covering 
(peridium) bears the hymenium, i.e. those elements from which 
the conidia are abstracted. The conidiocarps arise either imme- 
diately from the hyphoe or from a stroma in which they are 
generally embedded. Conidiocarps are entirely wanting in the 
Phycomycetes. On the other hand they are found among the 
Ascomycetes and Basidiomycetes, and in the latter group the 
conidiocarps contain more highly differentiated conidiophores 
(basidia) and are known as hasidiocarps. Conidiocarps with simple 
conidiophores, are found only among the Basidiomycetes, in the 
Uredinacese, and in Craterocolla cerasi. In the Ascomycetes (Figs. 
120 cZ, e ; 117 a, b ; 123 a ; 124? 6) the conidiocarps are visible, as 
points, to the naked eye, while the basidiocarps of the Basidiomy- 
cetes (Figs. 170, 171, 173-176, 178-180) vary from the size of a 
pea to that of a child's head. The " spermogonia " of the 
Ascomycetes and Lichenes, are conidiocarps with small conidia 
(microconidia) which germinate sometimes more slowly than other 
conidia, and formerly were erroneously considered as male repro- 
ductive cells, and called spermatia. 

The conidia of the Fungi are not primitive structures. The 
comparison of the sporangia and conidia among the Zygomycetes, 
and among the species of the genus Peronospora shows, that the 
conidia are aberrant formations, and that they have arisen through 

90 FUNGI. 

the degeneration of the sporangium, which, by the reduction of its 
spores to one, has itself become a spore. 

In the genera Thamnidium and Clxetocladium the gradual diminution of the 
sporangia, and the reduction of the number of spores can be distinctly followed. 
In Thamnidium the number of spores is often reduced to one, which is free in 
the sporangium. In Chatocladinm however the sporangia are typically one-spared, 
the spore is always united with the sporangium, and the two become a single 
body, the so-called conidium, which is in reality a closed sporangium. How 
close is the connection between the sporangia and conidia of Thamnidium 
and Chtetocladium, is seen from the fact that, in the conidial stage of 
Chcetocladium the same whorl-form of branching appears as in the sporaugial 
stages of Thamnidunn ch(etocladloides, and also, that the conidia of Ch. frexen- 
ianum throw off the former sporangium-wall (exosporium), while Ch. jonexii 
germinates without shedding its exosporium. The Phycomycetes have doubt- 
less sprung from Water-Alga? and inherit the sporangia from them. On this 
supposition, as the Phycomycetes assumed a terrestrial mode of life, the sporangia 
would become adapted to the distribution of the spores by means of the air, 
the sporangia would become small, contain dust-like spores, and would 
eventuall}' become closed-sporangia, i.e. conidia. The conidia are a terrestrial 
method for the multiplication of Fungi. In the Hemiasci and the Ascomycetes 
the sporangia are still preserved, but in every instance they are adapted to 
terrestrial spore-distribution, their spores being set free on the destruction of the 
sporangium-wall (generally shot out) and distributed through the air. For 
further examples of spore-distribution see below, p. 91-93. 

The reproduction of Fungi is accomplished not only by spores 
and conidia, but also sometimes by chlamydospores. These are 
fundaments * of sporangiophores and conidiophores, which have 
taken on a resting condition in the form of a spore, and are able 
to germinate and produce carpophores. In the formation of the 
chlamydospores the hyphre accumulate reserve materials at the 
expense of the neighbouring cells ; in the undivided hyphse of the 
Phycomycetes transverse walls are formed, and finally the chlamy- 
dospores are set free by the decay of the empty cells connecting 
them with the mycelium. One must distinguish between oidia and 
true chlamydospores. The former are more simple, the latter are 
a somewhat more differentiated form of carpophore fundaments, 
which serve for propagation in the same manner as spores. In 
CJilamydomucor racemosns the chlamydospores grow out into the 
air and form differentiated carpophores. In the Autobasidiomy- 
cetes they only gei'minate vegetatively, and not with the forma- 
tion of fructifications. From Cldamydomucor up to the Auto- 
basidiomycetes the successive development of the fructification, 

1 This term is adopted as a translation of the German " anlage." 

FUNGI. 91 

which is interrupted by the formation of the chlamydospores, 
degenerates more and more. Among certain Ustilaginere the 
chlamydospores (brand-spores) no longer germinate with the pro- 
duction of fructifications. In the Uredinacea?, only one of the three 
chlamydospore-forms has the property of producing fructifications 
on germination; the other forms only germinate vegetatively, like 
ordinary spores, and in the same manner as the chlamydospores 
of the Autobasidiornycetes. In the Hemibasidii, and the Uredin- 
acea?, in Protomyces, the chlamydospores are the chief means of 
reproduction. They are found also among the Ascomycetes. 

The sporangia and the conidia of the Fungi have their common 
origin in the sporangia of the Phycomycetes. The asci (and 
the Ascomycetes which are characterised by these bodies) are 
descended from the sporangia-forming, lower Fungi ; the basidia 
(and the Basidiornycetes) from those which bear conidia. The 
sporangia of the Phycomycetes are the primitive form and the start- 
ing point for all the reproductive forms of the Fungi. The chlamy- 
dospores appear besides in all classes of Fungi as supplementary 
forms of reproduction, and are of no importance in determining 
relationships. Although the expression "fruit" must essentially 
be applied to true Phanerogams, yet, through usage, the term 
" fruit- forms " is employed to designate the forms or means of 
reproduction of Fungi, and the organs of reproduction are known 
as organs of fructification, the sporangiophores and conidiophores 
as fruit -bearers (carpopliores), and the sporangiocarps, conidiocarps, 
and basidiocarps as "fruit-bodies." 

The majority of Fungi have more than one method of reproduction, often on 
various hosts (Uredinaceae). Species with one, two, or more than two methods of 
reproduction are spoken of as having monomorphic, dimorphic, or pleomorphic 
fructification. Monomorphic, e.g. the Tuberaceas ; dimorphic, Mttcor, Pipto- 
cepJialis, Saprolegniacea?, Penicillium crustaceum ; pleomorphic, Pucciiriu 
graminis, Capnodium salicinum (in the last species there are five methods of 
reproduction : yeast-like conidia, free conidiophores, conidiocarps with small 
and large conidia, and ascocarps). 

The liberation and distribution of the spores and 
conidia. The spores and conidia, on account of their small size 
and lightness, are spread far and wide by currents in the air, but 
in addition to this method, insects and other animals frequently 
assist in disseminating them. The liberation of the conidia is 
occasionally effected by the complete shrinking away of the 
conidiophore, but more frequently by abstriction from the conidio- 
phores, either by their gradually tapering to a point, or by the 

y-i FUNGI. 

dissolution of a cross-wall (generally of a mucilaginous nature). 
The individual links of conidia-chains are detached from one 
another in the same way, or often by means of small, intercalary 
cells, which are formed at the base of the individual links, and 
becoming slimy, dissolve upon the maturity of the spores. 
Special contrivances for ejecting the spores and conidia may often 
be found. In Peronospora the cylindrical fruit-hyphaa in the dry 
condition become strap-shaped and also twisted. These are very 
hygroscopic, and the changes of form take place so suddenly, that 
the spores are violently detached and shot away. In Empusa a 
peculiar squirting mechanism may be found (Fig. 85). Each club- 
shaped hypha which projects from the body of the fly, bears a 
conidium at its apex ; a vacuole, which grows gradually larger, 
is formed in the slimy contents of the hypha, and the pressure 
thereby eventually becomes so great that the hypha bursts at its 
apex, and the conidiam is shot into the air. By a similar mechan- 
ism, the spores of many of the Agaricaceae are cast away from 
the parent-plants. In the case of Pilobnlus (Fig. 84) the entire 
sporangium is thrown for some distance into the air by a similar 
contrivance, the basal region of the sporangium having, by the 
absorption of water, been transformed into a slimy layer which 
is readily detached. Sphcerobolus, a Gasteromycete, has a small, 
spherical fruit-body (basidiocarp), the covering of which, when 
ripe, suddenly bursts, and the basidiospores contained in it are 
forcibly ejected. 

The spores ichich are enclosed in asci are, in some instances, set 
free from the mother-cell (ascus) prior to their complete develop- 
ment (Elaphomycet, Enrotiuni). In the case of the majority of the 
Pyrenomycetes and Truffles, the asci swell by the absorption of 
water into a slimy mass, which gradually disappears, so that the 
spores lie free in the fruit-body ; they either remain there till the 
fruit- body decays, as in those which have no aperture (Perisporiaceae, 
Tuberaceae), or the slimy mass, by its growth, is forced out through 
the aperture of the sporocarp, taking the spores with it (Nectria). 
The ejection of the spores by mechanical means takes place in a 
number of Ascomycetes, and should many spores be simultane- 
ously ejected, a dust-cloud may be seen with the naked eye to 
arise in the air from the fruit-body. This is the case in the larger 
species of Pe:iza, Helvella, Rhytisma, when suddenly exposed to a 
damp current of air. A distinction is drawn between a simultaneous 
ejection of all the spores contained in the ascus, and an ejection at 

FUNGI. 93 

intervals (successive), when only one spore at a time is thrown out. 
The first of these methods is the most frequent, and is brought 
about by the ascus being lined with a layer of protoplasm, which 
absorbs water to such a degree that the elastic walls are extended 
at times to double their original size. The spores are forced up 
against the free end of the ascus, a circular rupture is made at 
this point, and the elastic walls contract, so that the fluid with 
the spores is ejected. Special means may in some instances be 
found to keep the spores together, and compel their simultaneous 
ejection. Thus, a tough slime may surround all the spores (Suc- 
cobolus), or a chain-apparatus, similarly formed of tough slime ; 
or there may be a hooked appendage from each end of the spores 
which hooks into the appendage of the next spore (Sordaria). The 
paraphyses occurring between the asci in many Ascomycetes, also 
play a part in the distribution of the spores, by reason of the 
pressure they exercise. The asci in some of the Pyrenomycetes, 
which are provided with jar-shaped fruit-bodies, elongate to such 
an extent that, without becoming detached from their bases, they 
reach the mouth of the fruit-body one at a time, burst and disperse 
their spores, and so make room for those succeeding. An ejection 
of the spores at intervals from the ascus is rarer. It takes place, 
for instance, in Pleospora, whose asci have a double wall. The 
external wall, by absorption of water, at last becomes ruptured, 
and the internal and more elastic membrane forces itself out in 
the course of a few seconds to one of two or three times greater 

, O 

length and thickness, so that one spore after another is forcibly 
ejected from a narrow aperture at the end of the ascus. 

Germination of spores (conidia and chlamydospores). In 
many spores may be found one or more germ-pores, i.e. thinner 
places, either in the inner membrane (uredospores, Sordaria) 
or in the external membrane (teleutospores in Rust-Fungi), 
through which the germination takes place. Generally this does 
not occur till the spores have been set free : in some Ascomycetes 
germination commences inside the ascus (Taphrina, Sclerotitiia ) . 
The different ways in which the spores germinate may be classified 
into three groups. 

I. THE ORDINARY GERMINATION occurs by the spore emitting a 
germ-tube, which immediately developes into a mycelium. In 
spores with a double wall it is only the inner membrane which 
forms the germ-tube. In swarmspores a single wall is formed 
after the withdrawal of the cilia, and this, by direct elongation, 

94 FONGl. 

becomes the germ-tube. The protoplasm accumulated in the 
spore enters the hypha, which, in pure water, can only grow as 
long as the reserve nourishment lasts. 

2. GERMINATION WITH PROMYCELIUM differs only by the circum- 
stance that the hypha developed from the germ-tube has a very 
limited growth, and hence it does not immediately develope into a 
mycelium, but produces coniclia (Rust- and Brand-Fungi). This 
promycelium must only be regarded as an advanced development 
of a conidiophore or basidium. 

3. THE YEAST-FORMATION of conidia consists in the production 
of outgrowths, very much constricted at their bases, from one or 
more places. Each of the conidia formed in this manner may 
again germinate in the same way. When sufficient nourishment is 
present, a branched chain of such conidia is formed, and these are 
finally detached from one another. Yeast-like buddings from the 
conidia are produced in various Fungi, e.g. Ascoidea, Protomyces, 
Ustilaginese, Ascomycetes, Tremellaceae, etc. In the Ustilaginese 
these conidia are an important element in the development. 
The budding conidia of Exobasidium forms a "mould" on the 
nutritive solution. The yeast-like conidia are not to be con- 
founded with the " Mucor-yeast " (comp. Mucoraceae). For Sac- 
charomyces see Appendix to the Fungi, page 176. 

In a compound spore (i.e. when a mass of spores are associated 
together) each spore germinates on its own account. There are 
sometimes, however, certain among them which do not germinate, 
but yield their contents to those which do. 

The length of time for which conidia can retain their power of 
germination is shortest (being only a few weeks) in those having 
thin walls and containing a large supply of water (Peronosporaeefe, 
Uredinacese). In many spores a resting period is absolutely neces- 
sary before they are able to germinate (resting spores). It has 
been observed in some spores and conidia, that the faculty of 
germinating may be preserved for several years if the conditions 
necessary for germination remain absent (Ustilaginese, ISurotium, 
Penicillium) . 

The optimum, minimum and maximum temperatures required 
for the germination of the spores has been decided in the case of 
a good many Fungi. A large portion of the most common Fungi 
have their optimum at 20C., minimum at 1 2C, maximum at 
40C. In the case of pathogenic Fungi the optimum is adapted to 
the temperature of the blood. Fungi living in manure, whose 


spores are often adapted to germinate in the alimentary canals of 
warm-blooded animals, have an optimum corresponding to the 
temperature of these animals, but with a little margin. 

Systematic Division. The lowest class of the Fungi is that 
of the PHYCOMYCETES, which have an unicellular mycelium, sexual 
and asexual reproduction, and have doubtless sprung from spo- 
rangia-bearing, lower Green Alga?. From the Phycomycetes (and 
certainly from the Zygomycetes) spring two well defined branches, 
each with numerous distinct species; to the one branch belong the 
HEMIASCI and the ASCOMYCETES, to the other the HEMIBASIDII and 
the BASIDIOMYCETKS. Ascomycetes and Basidiomycetes may be 
united under the title of MYCOMYCETES or HIGHER FUNGI. The 
Hemiasci and the Hemibasidii constitute the class of MESOMYCETES. 
The Hemiasci are an intermediate form between Zygomycetes and 
Ascomycetes ; the Hemibasidii a, similar group between the Zygo- 
mycetes and Basidiomycetes. Mesomycetesand Mycomyceteshave 
only asexual reproduction ; sexual reproduction is wanting. Their 
mycelium is multicellular. 

Up to the present time about 39,000 species have been described. 

Review of the divisions of the Fungi : 

ciass L Phycomycetes (Algal-Fungi). 

Sub-Class 1. Zygomycetes. 
Sub-Class 2. Oomycetes. 




class ii. Mesomycetes. 

Sub-Class l. Hemiasci. 

Sub-Class 2. Hemibasidii (Brand-Fungi). 

ciass in. Mycomycetes (Higher Fungi). 

Sub-Class 1. Ascomycetes. 
Series 1. Exoasci. 
Series 2. Carpoasci. 

Family 1. GYMNOASCALES. -\ 

Family 2. PEHISPORIALES. ? Angiocarpic Exoasci. 


Family 4. HYSTERIALES 1 TT T< 

* - Hemi-angiocarpic Exoasci. 

family 5. DISCOMYCETES. J 

Family 6. HELVELLALES. Gymnocarpic (?) Exoasci. 
Additional : ASCOLICHENES. Lichen-forming Ascomycetes. 


Sub-Class 2. Basidiomycetes. 

Series 1. Protobasidiomycetes. Partly gymnocarpic, partly 


Series 2. Autobasidiomycetes. 
Family 1. DACRYOMYCETES. Gymnocarpic. 
Family 2. HYIIENOJIYCETES. Piirtly gymuocarpic, partly hemi- 


Family 3. PHALLOIDE^E. Herniangiocarpic. 
Family 4. GASTEROMYCETES. Angiocarpic. 
Additional : BASIDIOLICHENES. Licben-foruiiug Basidiomycetes. 
Additional to the Fungi : FUNGI IMPERFECTI. Incompletely known 
(Saccharomyees, Oidium-loims, etc.). 

Class 1. Phycomycetes (Algal-Fungi). 1 

This group resembles Vaucher.ia and the other Siphoneae among 
the Algre. 

ORGANS OF NUTRITION. The mycelium is formed of a single cell, 
often thread-like and abundantly branched (Fig. 78). Vegetative 
propagation by chlamydospores and oidia. Asexual reproduction 
by endospores (sometimes sicarmspores) and conidia. Sexual re- 
production by conjugation of two hypha? as in the Conjugates, or 
by fertilisation of an egg-cell in an oogonium. On this account 
the class of the Phycomycetes is divided into two sub-classes : 


Sub-Class I. Zygomycetes. 

Sexual reproduction takes place by zygospores, which function 
as resting-spores, and arise in consequence of conjugation (Fig. 81); 
in the majority of species these are rarely found, and only under 
special conditions. The most common method of reproduction is by 
endospores, by acrogenous conidia, by chlamydospores, or by oidia. 
Swarmspores are wanting. Parasites and saprophytes (order 6 and 7). 
The zygospores are generally produced when the formation of 
sporangia has ceased; e.g. by the suppression of the sporangial- 
hyphre (Mucor wucedo), or by the diminution of oxygen; Pilobolus 
crystaliinus forms zygospores, when the sporangia are infected 
with saprophytic Piptocepltalis or Pleotraclielus. 

A. Asexual reproduction only by sporangia. 

Order 1. Mucoraceae. The spherical sporangia contain many 
spores. The zygospore is formed between two unicellular branches 

1 Also termed Water-Fungi (Wasserpilzeii). 



The unicellular mycelium (Fig. 78) of the Mucoracese branches 
abundantly, and lives, generally, as a saprophyte on all sorts of 
dead organic remains. Some of these Fungi are known. to be cap- 
able of producing alcoholic fermentation, in common with the Sac- 
charomyces. This applies especially to Chlamydomucor racemosus 
(Mucor racemosus}, when grown in a saccharine solution, and de- 
prived of oxygen; the mycelium, under such conditions, becomes 
divided by transverse walls into a large number of small cells. 

FIG. 78. Mucor m'iccdo. A mycelium which has sprung from one spore, whose position 
is marked by the * : a, b, c are three sporangia in different stages of development ; a is 
the youngest one, as yet only a short, thick, erect branch ; b is commencing to form a 
sporangium which is larger in c, but not yet separated from its stalk. 

Many of these swell out into spherical or club-shaped cells, and 
when detached from one another become chlamydospores, which 
abstrict new cells of similar nature (Fig. 79). These chlamydo- 
spores were formerly erroneously termed " mucor-yeast," but they 
must not be confounded with the yeast-conidia (page 94). They 
are shortened hypha?, and are not conidia of definite size, shape, 
and point of budding. Oidia are also found in Clilamydomucor. 
w. B. H 



The Mucoracere, in addition to the chlamydospores and oidia, 
have a more normal and ordinary method of reproduction ; viz., 
by spores which are formed without any sexual act. Mucor has 
round sporangia ; from the mycelium one or more long- branches, 
sometimes several centimetres in length, grow vertically into the 
air; the apex swells (Figs. 78, 80) into a sphere which soon 
becomes separated from its stalk by a transverse wall ; in the 

interior of this sphere (spor- 
angium) a number of spores are 
formed which eventually are set 
free by the rupture of the wall. 
The transverse wall protrudes 
into the sporangium and forms 
the well-known columella (Fig. 
80 d, e). The formation of 
spores takes place in various 
ways among the different 

SEXUAL REPRODUCTION by conjugation takes place in the follow- 
ing manner. The ends of two hyphse meet (Fig. 81) and become 
more or less club-shaped ; the ends of each of these are cut off 
by a cell- wall, and two new small cells (Fig. 81 A) are thus' 
formed, these coalesce and give rise to a new cell which becomes 
the very thick-walled zygote (zygospore), and germinates after 

FIG. 79. Chlamydospores of Chlamydo- 
mucor racemosus (x 375 times.) 

FIG. 80. Mucor ?ni<cedo : a a spore commencing to germinate ( x 300 timet-); 6 a germi- 
nating spore which has formed a germ-tube from each end (x 300 times) ; c the apex of a 
young sporangium before the formation of spores lias commenced ; the stalk is protruded 
in the sporangium in the form of a column : on the wall of the sporangium is found a very 
fine incrustation of lime in the form of thorn-like projections ; d a sporangium in which 
the formation of spores has commenced ; c a sporangium, the wall of which is ruptured, 
leaving a remnant attached to the base of the columella as a small collar. A few spores are 
seen still adhering to the columella. 


peviod of rest, producing a new hypha, which bears a sporangium 
(Fig. 81 E). 

Mucor mucedo, Pin- 
mould, resembles some- 
what in appearance 
Penicillium crustaceum 
and is found growing 
upon various organic 
materials (bread, jam, 
dung, etc.). 

Pilobolus (Figs. 83, 
84) grows on manure. 
Its sporangium (Fig. 84 
a") is formed during the 
night and by a peculiar 
mechanism (page 92) is 
shot away from the plant 
in the course of the day. 
This generally takes 
place in the summer, 
between eight and ten 
a.m. The sporangium 
is shot away to a height 
which may be 300 times 
greater than that of the 
plant itself, and by its 
stickiness it becomes 
attached to portions of 
plants, etc., which are 
in the vicinity. If these 
are eaten by animals, 
the spores pass into the 
alimentary canal and 
are later on, sometimes 
even in a germinating 
condition, passed out 
with the excrement, in 
which they form new 

Pliycomyces nitens 
(" Oil-mould ") is the 

FIGS. 81, 82. Mucor mucedo: A-C stages in the formation of 
the zygote ; D zygote; E germination of zygote : the exospore 
has burst, and the endospore grown into a hypha bearing 
a sporangium. 

largest of the Mould Fungi ; its sporangiophores may attain the height of 10-30 c. m. 

Order 2. Rhizopaceae. Ilhizopus nigricans (Mucor stolonifer) which lives 
on decaying fruits containing sugar, on bread, etc., has, at the base of the 
sporangiophores, tufts of rhizoids, i.e. hyphre, which function as organs of 
attachment. From these, " runners " are produced which in a similar manner 
. develope sporaugiophores and rhizoids. 

Order 3. Thamnidiaceas. On the same sporangiophore, in addition to a 



large, terminal, many-spored sporangium, many smaller, lateral sporangia are 
formed with a few spores. Thamnidium. 

B. Asexual reproduction by sporangia and conidia. 

Order 4. Choanephoraceae. Choanephora with creeping endophytic my- 
celium, and perpendicular sporangiophores. 

Order 5. Mortierellaceae. Mortierella polycephala produces on the same 
mycelium conidia and sporangiopbores. M. 
rostafinskii has a long stalked sporangiophore, 
which is surrounded at its hase by a covering of 
numerous felted hjphae. 

FIG. 83. Pilobolus. Mycelium (a, a), 
with a sporangiophore (A) and the 
fundament of another ( B). 


FIG. 84. Pilobolus. Sporangium (a") with 
stalk (a-c), which is covered by many 
small drops of water pressed out by tur- 

C. Asexual reproduction only by conidia. 

Order 6. Chaetocladiaceae. The conidia are abstricted singly and acro- 
genously. Chtetocladium is a parasite on the larger Mucoraceae. 

Order 7. Piptocephalidaceae. The conidia are formed acrogenously and 
in a series, by transverse divisions. The zygospore arises at the summit of the 
conjugating hypha?, which are curved so as to resemble a pair of tongs. Pipto- 
cephalis and Syncephalis live parasitically on the larger Mucoraceae. 

Sub-Class 2. Oomycetes. 

Sexual reproduction is oogamous with the formation of brown, 
thick-walled oospores which germinate after a period of rest. 
Asexual reproduction by conidia and sicarmspores. Parasites, 
seldom saprophytes. 

The oospores are large spores which are formed from the egg- 
cell (oosphere) of the oogonium (oosporangium, Fig. 89, 95). A 
branch of the mycelium attaches itself to the oogonium and forms 
at its apex the so-called " antlieridium " (pollinodium 1 ) : this sends 
one or more slender prolongations (fertilising tubes) through the 
wall of the oogonium to the egg-cell. 

1 Antheridium is preferred in this sub-class as keeping a more uniform term (Ku). 



A fertilisation, a passage of the contents of the anthcridium to the egg-cell, 
has as yet only been observed in Pythium; in Phytophthora only one small 
mass of protoplasm passes through the fertilising tube to the egg-cell ; in Pero- 

FIG. 85. Empusa muscit (Fly-mould). I. A fly killed by the fungus, surrounded by a whit 
layer of conidia. II. The conidiophores (t) projecting from the body of the fly. Some of 
the conidia, a few of which have developed secondary conidia, are attached to the hairs 
(mag. 80 times). III. A perfect hypha. IV. A hypha in the act of ejecting a conidium 
(c), enveloped in a sticky slime (9). V. A conidium which has developed a secondary 
conidium (sc). VI. A branched hypha produced by cultivation. VII. A secondary con- 
idium which has produced a small mycelium (in.)- VIII. A conidium germinating on the 
fly's body. IX, Mycelium. X. Conidia germinating like yeast in the fatty tissue of the 
fly. (III.-VII. and IX. magnified 300 times ; vm. and X. magnified 500 times.) 


nospora and the Saprolegniacefe'no protoplasm can be observed to pass through 
the fertilising tube, so that in these instances parthenogenesis takes place ; 
Saprolegnia tlmretii, etc., have generally even no antheridia, but nevertheless 
form normal oospores. Fertilisation of the egg-cell by means of self-motile 
spermatozoids is only found in Monoblepharis sphcerica. 

A. Asexual reproduction by conidia only. 

Family 1. Entomophthorales. 

The mycelium is richly branched. The family is a transitional 
step to the conidia-bearing Zygomycetes, since the oospores of many 
members of this family arise, and are formed, like zygospores. 

Order 1. Entomophthoraceae. Mycelium abundantly de- 
veloped. This most frequently lives parasitically in living insects, 
causing their death. The conidiophores forming the conidial- 
layer project from the skin, and abstrict a proportionately large 
conidium which is ejected with considerable force, and by this 
means transferred to other insects. These become infected by 
the entrance of the germ-tube into their bodies. The spherical, 
brown resting-spores develope inside the bodies of insects and 
germinate by emitting a germ-tube. 

GENERA : Empusa has a good many species which are parasitic on flies, 
moths, grasshoppers, plant-lice. The conidia emit a germ-tube which pierces 
the skin of the insect ; a number of secondary conidia are then produced inside 
its body, by division or by gemmation similar to that taking place in yeast, each 
of which grows and becomes a long unbranched hypha, and these eventually 
fill up the body of the animal, causing distension and death. Each of these 
hyphse projects through the skin, and abstricts a couidium, which is ejected 
by a squirting contrivance. The best known species is E. muscle (Fig. 85), 
which makes its appearance epidemically towards autumn on the common 
house-fly, and shows itself by the dead flies which are found on the windows 
and walls attached by their probosces, distended wings, and legs. They have 
swollen abdomen, broad white belts of hyphje between the abdominal rings, and 
are surrounded by a circle of whitish dust formed by the ejected conidia. 
Entomophthora sends out, at definite places, from the mycelium hidden in the 
insect's body, bundles of hyphas, which serve the purpose of holding fast the 
dead insects, the ramifications attaching themselves to the substratum : the 
conidiophores are branched, the conidia are ejected by the divisional walls be- 
tween the hyphre and the conidia dividing into two layers, those which ter- 
minate the hyphae suddenly expanding and throwing the conidia into the air. 
E. radicans makes its appearance epidemically on caterpillars. 

B. Asexual reproduction by zoospores or conidia. 

Family 2. Chytridiales. 

In this family the mycelium is very sparsely developed or is 
wanting. The entire plant consists principally or entirely of a 



single zoosporangium. whose zoospores have generally one cilium. 
The resting-spores arise either directly from the zoosporangium, 
which, instead of forming zoospores, surrounds itself by a thick 
cell-wall; or they are formed by the conjugation of two cells (in 
which case they are spoken of as oospores). Microscopic Fungi, 
parasitic on water plants (especially Algae) or small aquatic ani- 
mals, seldom on land plants. 

Order 1. Olpidiaceae. Without mycelium. Swarmspores 
and resting-spores. 

Iu the Olpidieie, the swarmspores, probably, most frequently form themselves 
into a plasmodium (naked mass of protoplasm) which may become a single 
zoosporangium or a resting sporangium. Olpidium trifolii occurs in Trifoliuni 
repens. In the Synchytriece the plasmodium emerging from the swarmspores 
breaks up either at once, or after a period of rest, into smaller plasmodia, each 
of which will become a zoosporaugium. Synchytrium anemones is found on 
Anemone nemorosa ; 8. mercuriulis on Mercurialis perennis ; S. aureum on many 
plants, particularly Lysimachia nummularia. 

FIG. 86. Ghytridium lagcnuln. Zoo- 
sporangium a before, b after the libera- 
tion of the swarmspores. 

FIG. 87. Obelidium mucronatum, : 
m mycelium ; s swarmspores. 

Order 2. Rhizidiaceae. Mycelium present. Zoospores and 

Chytridium (Fig. 86). Obelidium (Fig. 87) is bicellular; the one cell is the 
mycelium, the other the zoosporangium ; found on insects. The species of 
Cladochytrium are intercellular parasites on marsh plants. Physoderma. 

Order 3. Zygochytriaceae. Mycelium present. Zoospores 
and oospores. The latter are the product of the conjugation of 
two cells (Fig. 88). 

Polyphayus euglmuc on Euglena viridis. Urophlyctis pulposa on species of 



Family 3. Mycosiphonales. 

The mycelium is bladder-like or branched. Zoospores. Sexual 
reproduction by oospores, which are produced in oogonia. The 
latter are fertilised, in some forms, by the antheridium. 

Order 1. Ancylistaceae. The entire bladder-like mycelium is used for the 
construction of zoosporangia, oogonia, or antheridia. Lagenedium is parasitic 
on Spirogyra, etc. 

Order 2. Peronosporaceae. Almost entirely parasites. The 
unicellular, often very long and abundantly branched mycelium 
lives in the intercellular spaces of living plants, especially in 
the green portions, and these are more or less destroyed and de- 
formed in consequence. Special small branches (suction-organs, 

FIG. 88. Polyphagits cuglcna:. A -with smooth, S with thorny oosporcs ; m and /the two 
conjugating cells. 

" Jiaustoria ") are pushed into the cells in order to abstract 
nourishment from them. Both oospores and conidia germinate 
either immediately, or they develope into sporangia with swarm- 
spores, having always two cilia. Only one oospore is formed in 
each oogonium ; its contents (Fig. 89) divide into a centrally 
placed egg-cell and the " periplasm " surrounding it; this is of a 
paler colour and on the maturity of the oospore forms its thick, 
brown, external covering. 

The Potato-fungus (PhytopJtthora infestans) is of great interest. 
Its thallus winters in the Potato-tuber; other organs for pass- 
ing the winter, such as oospores, are not known. AVhen the tuber 
germinates, the Fungus-hyphre penetrate the young shoot and 
keep pace with the aerial growth and development of the plant. 
The conidiophores emerge through the stomata, especially on the 
under side of the leaves ; they branch like a tree (Fig. 90), and 



appear to the naked eye as a fine mould on the surface of the 
plant. The disease soon makes itself known by the brown colour- 
ing of those parts of the plant which are attacked, and by their 
withering. An ovoid conidium arises at first by the formation of 
a dividing wall at the apex of each branch of the conidiophove, 

FIG. 89. Peronospora. alsinearum. My 
celium with egg-cell and antheridium. 

FIG. 90. Ph\jtoplithora infestans (strongly magnified). Cross section through a small 
portion of a Potato-leaf (the under side turned upwards): a the mycelium ; b b two conidio- 
phores projecting through a stoma; c conidia; e the spongy tissue of the leaf; g the 



FIG. 91. PhytopTithora infestans: n-c conidia de- 
tached ; in c the swarm-cells are leaving the mother- 
cell; d two free-swimming swarm-cells. 

(Fig. 90 cc), and immediately underneath it another is formed, 
which pushes the first to one side, and so on. These conidia some- 
times germinate directly, and form a mycelium, but most frequently 
their protoplasm divides into many small masses, each of which 
becomes a pear-shaped zoospore provided with two cilia (Fig. 91). 
Water is required for their germination, and when the ripe conidia 

are placed in a drop of 
water the swarm-cells are 
formed in the course of 
about five hours. They 
swarm about in rain and 
dewdrops in the Potato- 
fields, and are carried with 
the water to the Potato- 
plants and to the tubers 
in the soil. The wind also 
very easily conveys the conidia to healthy Potato-fields and infects 
them. The enormous quantity of conidia and swarm-cells that 
may be formed in the course of a summer explains the rapid 
spreading of the disease ; and the preceding makes it clear why 
wet summers are favourable to its existence. When the swarm- 
cells germinate, they round off, and then surround themselves 
with a cell-wall which grows out into the germ-tube, and pierces 
through the epidermis of the host-plant (Fig. 92). Having entered 

the host, a new mycelium 
is formed. The potato 
disease, since 1845, has 
been rampant in Europe ; 
it has, no doubt, been in- 
troduced from America, 
which, it must be remem- 
bered, is the home of the 

FIG. 92. Pltytophthora infestans. Cross section 
through a portion of a Potato-stalk. Two germinal- 
ing conidia (a, b) piercing the epidermis, and the 
mycelium penetrating the cells. 

The conidia exhibit -various 
characters winch are employed 
for the separation of the genera. 
Pythium is the most simple 
form. The contents of the ter- 
minally-formed conidia emerge as a spherical mass and divide into swarmspores. 
P. de Ilaryanum lives in the seedlings of many different Flowering-plants, 
which it completely destroys. Phytophtlwra is distinguished by the circum- 
stance that the sparsely-branched couidiophores bear, syinpodially, chains of 



conidia. Besides the Potato-fungus (see above), Ph.fagl belongs to this group ; 
it developes oospores very abundantly, and does great harm to seedlings of the 
Beech, Sycamore, and Pine trees. Peronospora generally has conidiophores 
which are repeatedly forked, and bear a conidium on each of the most extreme 
ramifications. Many do great harm to their host-plants. P. viticola, on Vines, 
and P. nivea, on umbelliferous plants, have swarmspores, which are absent in 
the following species of this genus : P. sparser, on Roses ; P. gangliformis, on 
composites ; P. alsinearum, on Stitchwort ; P. parasitica, on cruciferous plants ; 
P. viciee, on Vetches and Peas ; P. schachtii, on Beets ; P. violacea, on the 

FIG. 93. A fly overgrown 
with Sctprolegnia, 

FIG. 94. Formation of swarmspores in a Saprolegnia : 
a germinating swarmspores. 

flowers of Scaliosa ; P. radii, on the ray-florets of Matricaria. Cystopus (Albugo) 
has the conidia developed in chains, which form a cohesive white layer under- 
neath the epidermis of the host-plant. Cy.itnpus Candidas, on cruciferous plants, 
especially Shepherd's Purse and Brassica ; the germination commences on the 
cotyledons, and from this point the mycelium developes together with the host- 
plant ; C. cubicus, on the leaves of Composite. 

Order 3. Saprolegniaceae, Water-Fungi which live as sapro- 
phytes on organic remains lying in water, for instance, on dead 
flies (Fig. 93), worms, remains of plants ; but they may also make 
their appearance on living animals, being 
frequently found, for example, on the 
young trout in rearing establishments. 

The thalhis is a single, long and branched 
cell. It has one portion which serves as 
root, and lives in the substratum, where it 
ramifies abundantly for the purpose of ab- 
sorbing nourishment ; and another portion 
projecting freely in the water, and sending 
out hyphee on all sides (Fig. 93). The 
asexual reproduction takes place by swarm- 
spores (Fig. 94), which are developed in 
large sporangia; these swarmspores gener- 
ally possess two cilia, and on germination FlG - 6--Oogonium with 

two antheridia, Achlya race- 
grow into new plants. The entire proto- -mesa. 


plasm in the oogonium is formed into one or more oospheres, 
Avithout any surrounding " periplasm." The oospheres may not 
be fertilised (p. 100), and then develope parthenogenetically. 

Genera : Saprolegnia, whose swarmspores disperse immediately after having 
left the sporangium. S. ferax is the cause of a disease in fish ("Salmon 
disease ") and in the crayfish. Achlya, whose swarmspores accumulate in a 
hollow ball before the mouth of the sporangium. Leptomitus has strongly in- 
dented hypha?, causing a " linked " appearance. L. lacteus is frequent in the 
waste matter from sugar factories. Monoblepharis deviates from the others by 
the greater development of its fertilising process ; the oosphere, situated in an 
open oogonium, becoming fertilised by self-motile spermatozoids, which are 
provided with a cilium at the posterior end. 

Class 2. Mesomycetes. 

The Mesomycetes are intermediate forms between the Phy- 
comycetes and the Higher Fungi. In the vegetative organs, and in 
the multicellular hyphse, they resemble the Higher Fungi; th 
methods of reproduction, however, show the characters of the Phy- 
comycetes, namely sporangia and conidiophores of varying size 
and with varying number of spores ; definite and typically formed 
asci and basidia are not present. Sexual repi-oduction is wanting. 
The HEMIASCI are transitional between the Phycomycetes and the 
Ascomycetes, the HEMIBASIUII (Brand-Fungi) form the transition 
to the Basidiomycetes. 

Sub-Class 1. Hemiasci. 

The Hemiasci are Fungi with sporangia which, although resem- 
bling asci, yet have not, however, a definite form and a definite 
number of spores. Besides endospores, conidia, chlamydospores 
and oidia are found. 

Order 1. Ascoideaceae. Ascoidea rubescens forms irregular, reddish-brown 
masses in the sap issuing from felled Beeches. It has free sporangia, which 
resemble asci in their structure, in the development and ejection, and in the 
definite shape and size of the spores. The formation of the sporangia takes 
place when the nutriment is nearly exhausted, and resembles that of the conidia, 
since they are developed from the end of a hypha which enlarges, and the swelling 
becomes separated by a transverse wall. Within the sporangia numerous spores of 
a cap-like form are developed, which are set free through an opening at the apex. 
Sporangia are formed successively at the apex of the same hypha, the second 
commencing to develope as the first is dehiscing. Conidia and sporangia are not 
formed simultaneously ; the former may be considered as closed sporangia. 

Order 2. Protomycetaceae. Protomyces pacJnjdcrmus causes hard swellings 
on the stems and leaf-stalks of the Cichorieaj (Taraxacum, etc.). These swell- 



ings consist of cJdamydospores (resting-spores), which germinate aud hecome 
free, ascus-like sporangia, with numerous small spores. In nutritive solutions 
the chlamydospores form conidia with yeast-like buddings. P. macro*porus on 
jEtjopodium, and other Umbelliferae. 

Order 3. Thelebolaceae. Thclebolus stercoreus, is found on the dung of 
deer, hares, and rabbits, and has clostd sporangia, which resemble asci in their 
shape and regular construction, and in the ejection of spores. The covering 
encloses only one sporangium, even where the sporangia arise close together. 

This order, bj reason of the covering 1 of the sporangia, forms the 
transition from the Hemiasci to the Carpoasci, while the two first 
supply an intermediate step to the Exoasci. 

Sub-Class 2. Hemibasidii, Brand-Fungi. 

The Brand-Fungi (also known as USTILAGINE^:) are Fungi with 
basidia-like conidiophores, which, however, have not yet advanced to 
a definite form or number of conidia. They are true parasites, 
whose mycelium spreads itself in the intercellular spaces of Flower- 

PIG. 96. Entyloma ranunculi. 1. Cross section of a portion of a leaf of Ficarin. permeated 
by the mycelium ; a bundle of hyphte with conidia emerging from a stoma ; in oue of the 
cells are found four brand-spores. ~2. A brand-spore developed in the middle of a hypha. 

ing plants. The mycelium is colourless, quickly perishable, has 
transverse walls at some distance from each other (Fig. 96), and 
sends out haustoria into the cells of the host-plant. 

It most frequently happens that the germ-tube enters the host- 
plant at its most tender age, that is, during the germination of the 
seed ; the mycelium then wanders about in the tissues of the shoot 



chiving its growth, until it reaches that part of the plant where 
the spores are to be formed. The spore- formation takes place in 
the same way in all those species whose brand-spores are deve- 
loped in the floral parts of the host-plant. Many Brand-Fungi have, 
however, a more local occurrence, and the mycelium is restricted 
to a smaller area of the leaf or stem. Those organs of the host- 
plant in which the brand-spores are developed often become 
strongly hypertrophied. In perennial plants the mycelium winters 
very often in the rhizome. 

The brand-spores are the winter resting-spores of the Brand- 
Fungi. They arise in the tissues of the host-plant, which is often 
destroyed, and become free through the rupture of the epidermis ; 

FIG. 97. Doassansia alismatis. 1. A fruit-body, formed by a covering of oblong 1 hypba% 
which encloses a mass of brand-spores, and is embedded in the leaf-tissue of the host- 
plant; 20 times natural size. 2. A germinating brand-spore, 500 times natural size. 3. 
Three connected resting-spores, 400 times natural size. 4. Two conidia grown together, 
600 times natural size. 

they are thick-walled, generally brown or violet, and very often 
possess warts, spines, or reticulate markings. Fruit-bodies, that is 
enclosed organs of reproduction, are found in few genera (Sphace- 
lotheca, Graphiola ; Doassansia, Fig. 97). In Tolyposporium, Tiibur- 
cinia, Thecapliora (Fig. 102), etc., the brand-spores are united into 
a ball of spores. On germination the brand-spores behave as 
cMamydospores, namely, as the fundament of conidiophores, by 
emitting a short germ-tube, i.e. a conidiophore (" promycelium"). 
The USTILAGINACE.E (Fig. 99, 2) have a short transversely divided co- 
uidiophore, with laterally developed conidia (comp. the basidia of 
the Protobasidiomycetes). The conidiophores of the TILLETIACE^: 
are iindivided (unicellular promycejia), and bea,r the conidia ter- 
minally, and so resemble the basidia of the Autobasidiomycetes. 



In Tilletia, Entyloma, Neovossia, Tuburcinia, the brand-spores germinate 
and form basidia-like conidiophores with spindle-shaped conidia ; their niy- 

FIG. 98. Tuburcintn. 1. T. trientalis. Hyphse, some of which bear conidia at the apex, 
forcing themselves out between the ep-'dermal cells on the under side of the leaf ; 320 times 
natural sue. 2. T. trientalis. A ball of spores in which some of the individual brand- 
spores are about to germinate ; 520 times natural size. 3. T. primulicola : various forms 
of conidia (500 times natural size). 

celium, on the other hand, produces later only single, sickle-shaped conidia, so 
that two kinds of conidia are found, as in a few Basidiomycetes. In some 
species, e.g. Ustilago hordei, the 
brand-spores only germinate 
vegetatively and form a myce- 
lium. In nutritive solutions (so- 
lutions of dung, etc.) where they 
live as sapropltytes, the brand- 
spores of many species emit 
germ-tubes, and on these, yeast- 
like conidia are produced by 
repeated budding, which grow 
into mycelia only when tbe 
nutritive solution is exhausted. 
These conidia have not the 
power of producing alcoholic 
fermentation. The very numer- 
ous conidia, which are found 
in the dung of herbivorous ani- 
mals, are probably the yeast- 
conidia of Brand-Fungi. The 
brand-spores, which are eaten 
by animals with the grain and 
hay, pass into the dung and 
without doubt give rise to a 
very rich multiplication of 

The couidia (also called 

FIG. 99. Vftilago. 1. Formation of brand- 
spores. 2. Germinating brand-spore of U. peren- 
nans. 3. Germinating brand-spore of 17. cardui 
(after Brefeld). 4. U. fdiformis. a A brand-spore 
with developed basidium ; b another, with a coni- 
dium ; c with two conidia ; d with two conidia 
placed diametrically opposite to each other ; e, de- 
tached conidia which are growing into hyphse. 



"sporidia") of many species unite generally into an H-form (Figs. 97, 4; 
100 h ; 101, 4). This union in pairs does not, however, take place with a view 

FIG. 100. TiUctia tritici : a an ear of Wheat in which all the grains are attacked by Stink- 
brand ; b a blighted corn surrounded by the chaff ; c a blighted corn grown together witli a 
stamen; d the same cutacross ; c a brand-spore; /. g, h germinating brand-spores ; i germi- 
nating conidia ; j the mycelium ; fc-fc brand-spore-f'orming mycelium-threads, (c-li magni- 
fied iOO times; i-k 300 times.) 



to germination, there is no fusion of nuclei, and therefore in this " fusion " there 
is no sexual act. 

Order 1. Ustilaginacese. Conidiophores with transverse walls and lateral 
conidia. Ustitago (Fig. 99) generally developes its spores in the floral organs 
of its host-plant, the ovary or anthers, where they arise from hyphaa, and form 
a slimy mass which when mature becomes a black dust. 

To this order belong U. avena;, parasitic on Oats, U. hordei and U. nuda 
(U. jenseni), on Barley; these are the usual cause of " Smut" on cereals. U. 
hypodytes on straw of Elymus and Agropyrum. U. filiformis in the leaves of 
Glycerin. U. caricis transforms the fruits of various species of Carex into 
black, dusty balls. U. violacea developes its violet spore-powder in the anthers 
of the Caryophyllacete. U. tragopogonis, transforms entire inflorescences of 
Tragopogon into a black-violet mass. Among the largest are U. grandis, which 
causes the large swollen nodes in the stem of Pliragmitcs, and the Maize Blight, 
U. maydis, which produces outgrowths about the size of a hand on the spadix of 
the Maize. 

Order 2. Tilletiaceae. Conidiophores undivided, generally several conidia 
arise at their apices. Tilletia tritici, the Stinkbrand on Wheat (Fig. 100). The 
mycelium lives in Wheat-plants, producing its spores in the ovary after the 
whole interior of this body has been destroyed by the mycelium, with the ex- 
ception of the external layer of the wall of the ovary, which remains essentially 
unaltered and encloses the closely packed, firm mass of spores (Fig. 100 d). 
The grains of Wheat thus attacked are shorter and thicker than the sound ones, 
and the ears show the presence of this Fungus by their erect position, and 
the wide separation of the chaff (Fig. 100 ). The unpleasant odour of the 
ovary prior to the ripening of the spores, has given the name " Stinkbraud," 
and, in like manner, its hardness when it encloses the ripe spores, is the reason 
of its being also called '' Stonebrand." On account of this hardness, the dis- 
eased grains are readily harvested together with the healthy ones, which become 
infected by the spores at the threshing. 2'. lacvis (T. fattens) also occurs on 
Wheat and has smooth brand-spores. 

Entyloma (Fi^. 96), a genus with numerous species, which appear in 
spots on the leaves of the host-plant, and Tulurcinia (Fig. 98), which makes 
its appearance on the Primu- 
laceffi, produce white conidia- 
spots on the surface of the 
host-plant. The first-named 
has single spores, the latter 
has its spores closely massed 
together. Urocystis (Fig.101) 
has its spores surrounded by 
a number of small and lighter 
coloured barren spores. U. 
occulta, Rye-stem Blight, 
forms its spores in long 
streaks in the stems and 
leaves of the Eye, and does 
considerable damage. U. ce- 

FIG. 101. Urocystis. 1, U. coralloides. A spore-ball, 
magnified 450 times. 2-4, U. anemones: 2-3, brand- 
spores which are about to germinate (magnified 450 
times). 4, Conidia, the two in a state of fusion, a 
third with vacuoles and division-vrall, magnified 500 

W. R. 



piila; on Onions. U. viola forms large dark-violet swellings in the leaf-stalk 
and stems of Violets. Thecaphora (Fig. 102) appears in seedlings of Con- 
volvulus and Astragalus. 

As a means of protection against the Smut-Fungi which make their appear- 
ance on the different cereals, a submersion of the grains in a solution of blue 
vitriol (J%) for twelve hours, or better still, submerging for five minutes in 
water heated to 53-55 C (Jensen's method) is employed. 

FIG. 102. Thecapliora. 1, T. convolvuli, a ball of spore?, one of the brand-spores has emitted 
a septate branched conidiophore ( x 520). 2, T. nffniis, a ball of spores (x 520). 

Class 3. Mycomycetes, Higher Fungi. 

The MYCOMYCETES are not entirely aquatic in habit ; they have 
hyphae with transverse watts, but no sexual reproductive organs. 
The asexual reproduction takes place in very different ways ; by 
endospores (in asci), conidia, basidiospores, chlamydospores, and 
oidia. Swarmspores are never found. 

Two chief methods of reproduction may be distinguished, and 
hence the class may be divided into two large sub-classes : the 
ASCOMYCETES (with asci), and the BASIDIOMYCETES (with basidia). 

Sub-Class 1. Ascomycetes. 

The main characteristic which distinguishes the Ascomycetes is 
the ascus ; a name given to a sporangium of a definite shape and 
size, and containing a definite number of spores. The shape is 



generally club-like or spherical, the number of spores 8 (in some 
2, 4, 16 or more), see Figs. 103, 105, 108, 110, 113, 116, 120, 121, 
123, 129. 

In the lowest forms, the EXOASCI, the ascus springs directly from 
the mycelium without the formation of a fruit-body (i.e. asco- 
carp). In the higher forms, which contain many species, the 
CABPOASCI, the asci are united and form ascocarps which may be 
more or less enclosed (angiocarpic, hemiangiocarpic, and probably 

FIG. 103. Endogenous formation of spores in Peziza confluent. In the youngest asci 
there is only one nucleus (b, e) ; this divides into two (/) ; and the division is repeated so 
that there are 4 nuclei in c and 8 in g. These surround themselves with protoplasm and a 
cell-wall (h, i)- The protoplasm of the mother-cell is not entirely used up. 

The hyphfe of the Mycelium in some remain free, in others 
they are felted together and form thick strands or flat, cushion-like 
bodies (compare in particular the stromata of the Pyrenomycetes). 
Some species form sclerotia (Figs. 116, 128). 

Asexual reproduction by means of conidia is known in many 
species as the principal means of reproduction, and the one 
\vhich affords the most rapid means of distribution. The conidia 
may be produced on conidiophores (Fig. 109), in conidial-layers 
(Fig. 122), and often in conidiocarps (pycnidia, Figs. 120 d, e ; 


123 a ; 124 &.). These last occur partly as the so-called " spermo- 
gonia " (that is, pycnidia with microconidia). The conidiophores 
never approach the basidia. 

In many species the ascospores germinate and form conidia immediately 
(Nectria cinnabarina, Sclerotinia, Taphrina, eic.)j sometimes while they are 
still in the ascus and before their ejection (Taphrina, Fig. 105 a). In many in- 
stances the conidia by means of continued budding can, for a longer or shorter 
time, produce yeast-conidia, e.g. Taphrina. In many other cases the conidia 
arise from the germ-tubes of the ascospores, or at any part of the mycelium. 
The unripe asci of Taphrina, when placed in water, develope conidia at their 
apices. The Sclerotinia-species produce numerous conidia whose germination 
has never been observed. The formation of conidia and asci sometimes takes 
place on the same fruit-body. In Heterosphteria patella the conidia and asci 
are developed successively in the same fruit-body ; in the ascocarps of Derma - 
tea frangida and Sclerotinea sclerotiorum the formation of conidia may take 
place. The ascocarps frequently arise from the conidial-layers (Nectria cin- 
nabarina, etc.). This relationship of the two forms of reproduction to each 
other may be explained by considering that both have descended phylogene- 
tically from sporangia. 

Sometimes chlamydospores and oidia also appear in the Ascomy- 
cetes ; on germination, however, they do not, as in Protomyces, 
form sporangia, and on this account cannot be distinctly distin- 
guished from conidia. 

The asci are morphologically the highest form of reproduction 
and are always found at the close of the development of these 
Fungi ; the accessory forms of reproduction are first developed, but 
a well-defined alternation of generations does not occur. 

In the Ascomycetes there are more than 11,000 described species, which can 
be classed as follows : 
Series 1. EXOASCI. Only one order. 

Family 1. Gymnoascales, -\ 

2. Perisporiales, > Angiocarpic Carpoasci. 
,, 3. Pyrenomycetes, ^ 

4. Hysteriales, | Hemiangiocarpic Carpoasci. 
,, 5. Discoinycetes, ) 

,, 6. Helvellales, Gynmocarpic (?) Carpoasci. 
Additional Ascolicfienes : Lichen-forming Ascomycetes. 

Series 1. Exoasci. 

Ascomycetes with FREE ASCI; sometimes also conidia, chlamydo- 
spores and oidia. One order. 

Order. Taphrinacese. Of the genera belonging to this order, 
Taphrina, Endomyces, and Ascocorticium, the first is most important. 



Endomyces decipiens is a parasite in the fruit-body of Armillaria mellea ; 
E. magnusii lives in the gelatinous, fermenting exudations of Oak-trees ; Asco- 
corticium albidum is found under the bark of the Fir-tree. Endomyces has 
chlamydospores and oidia. 

The species of Taphrina, are parasites, whose free asci may be 
found in great numbers, generally closely pressed together, on 
the parts of plants which they have attacked. The asci are de- 
veloped directly from the ascogenous cells of a fertile, generally 
sub-cuticular, hypha, which arises from the sterile mycelium. The 
latter arises from the germinating ascospore, and may hibernate 
in the tissues of its host, particularly in the winter buds, and then 
with the commencement of the next period of vegetation it con- 


FIG. 104. Taphrtiia (Exoascus) 
pi-uni. Yeast-like budding of a 
germinating spore ( x 600). 


O Oo 



FIG. 105. Taplirina betuKna : a ascus filled 
with conidia; b germinating spores (x600). 

tinues its growth side by side with that of its host. The hyphae 
ramify in the intercellular spaces or beneath the cuticle, but have 
no haustoria. The ascospores (fig. 105 A) and unripe asci may 
produce conidia. 

Very remarkable appearances, and swellings of the attacked tissues, are pro- 
duced when the mycelium is perennial; for example, the " Witches' -brooms " 
and " Pockets." The hard, hollow, stoneless plums, known as " Pocket " or 
" Bladder " Plums, are produced by considerable changes in the tissues of the 
fruit ; these are caused particularly by T. pruni on several species of Prumis. 
The " Witches'-brooms," on the contrary, are deformations of entire twigs or 
branches, and often attain a very large size. They occur on Almis incana, 
caused by T. epiphylla ; on Carpinus betulus, by T. carpini ; on Cherry-trees, 
by T. cerasi ; on Plum-trees, by T. insitituc ; on Birches, by T. titryida and 
T. betulina. T. deformans attacks the leaves of the Peach, and causes them 
to curl. 


When a perennial mycelium is wanting, the infection is confined as a rule to 
white or yellow spots on the leaves, e.g. the commonest, T. sadebeckii, on Alnus 
glutinoia, and T. aurea on species of Populus. T. alni incance (Fig. 106) causes 
considerable hypertrophies on the pistillate catkins of the Alder, which may be 
compared to the " pockets " of Pninus. 

FIG. 106. Taphrina alni incance on the Alder (nat. size). 

Series 2. Carpoasci. 

The Carpoasci are Ascomycetes, whose asci are enclosed in fruit- 
bodies, i.e. ascocarps. The accessory means of reproduction are 
free conidiophores (Fig. 109), conidial-layers (Fig 1 . 122), conidio- 
carps (Fig. 120 _D, E, etc.), chlamydospores and oidia. 

For the different methods of distributing the ascospores, see p. 

Of the six families of the Carpoasci, the first three Gymnoa- 
scales, Perisporiales, and Pyrenomycetes are ANGIOCARPIC (that is, 
the ascocarp remains closed throughout its existence, and does 
not dehisce when ripe) ; the fourth and fifth families (Hysteriales 
and Discomycetes), on the other hand, are HEMIANGIOCARPIC (the 
ascocarp, here also called an apothecium, is closed in the early 
stages, but opens at the commencement of ripening and exposes 
a hymenium of crowded asci) ; the family of Helvellales has 
probably GYMXOCARPIC (or hemiangiocarpic) fruit-bodies. 

Family 1. Gymnoascales. 

The ascocarps are surrounded by a spongy and incomplete en- 
velope. One order, poor in species. 



Order Gymnoascaceae. The ascocarps are borne sometimes solitarily, or 
sometimes coiled together. Gymnoascus reessii forms small bodies about 1 
mm. in diameter on old horse-dung, which at first are white and afterwards 
orange-red. Ctenomyces serratus lives on the old feathers in birds' nests. 

Family 2. Perisporiales. 

The ascocarps are surrounded by a complete envelope without any 
opening : the fruit-bodies are cleistocarpic ; the spores are only 
liberated after the disintegration of the fruit-bodies. Paraphyses 
are wanting. The two first orders have in addition the means 
of reproduction by conidia. 

Order 1. Erysiphaceae, Mildews. The Fungi belonging 
to this order are epiphytic parasites, whose mycelium, somewhat 
resembling a cobweb, may be seen on the leaves and other 
green portions of plants (see Figs. 107, 108). The hyphte ramify 

FIG. 107. Erysiphe ciehoriacearum : a mycelium-threads; b sterile hypha ("pollino- 
clium") ; c fertile hypha (ascogone or archicarp) ; d and e young ascocarps. 

in all directions upon the surface of their host, and emit haustoria 
which penetrate the epidermal cells, and thus derive the necessary 
nutriment. The Mildew-Fungi thus belong to the obligate para- 
sites, and during their growth dwarf and destroy the portions 
of their host on which they live. The reproduction takes place in 
the first instance by abstriction of conidio-chains from the end of 
special branches (Fig. 108 c, a hypha is seen in the act of detaching 
a conidium). The conidia may germinate immediately, and thus 
quickly reproduce their species. When present in large numbers 
they appear as a white meal covering the surface of the plant on 
which the fungus is found. Later on appear the dark brown, 
spheroid ascocarps (Fig. 108 a) which, although small, are gene- 
rally just visible to the naked eye as black specks. 

A characteristic feature of the Mildew-Fungi is the thin, pseudo- 



parenchymatons covering 1 of the ascocarp, enclosing one (Podo- 
sphcera and Sphrerotheca ; compare Thelebolus among the Hemi- 
asci) or a few asci (Fig. 108 c), which do not form any hymenium, 
but are irregularly placed. The cells of the ascocarp-envelope 
are often prolonged into hair-like appendages. The ascocarps are 
developed from the mycelium at places where two hyphre ci-oss 
each other (Fig. 107). At these places two short and erect hyphte 
are produced side by side. The one from the lower hypha (Fig. 
107 o) assumes an ellipsoidal shape, and is known as the arclticarp 
or ascogone, while the other (" pollinodium ") arches over the 

FIG. 103. Ei-ysiphe communis. A small portion of a leaf with this Fungus growing upon 
it (considerably magnified). The hyphse 6 and d do not belong to this Fungus, but are re- 
productive organs of a pyrenomycetons Fungus parasitic upon it (Cicinnobotus). 


From the latter one ascus may be at once developed 
(Sphcerotheca, etc.), or after its division several asci may be pro- 
duced, each developed from one division. The sterile hypha 
(termed " pollinodium," since it was formerly, but erroneously, 
supposed to fertilise the ascogone) produces a number of branches, 
and forms the pseudo-parenchymatous envelope of one cell in 
thickness, enclosing the asci. 

Many plants, both cultivated and wild, are attacked by various 



species of Mildew. A common means of prevention against their 
attacks is to dust the diseased parts with sulphur. 

Spharotlieca pannosa occurs on the leaves of Roses, and on the fruit of 
Peaches and Apricots. S. castagnei on Humiilus, Cucitmis, etc. Erysiplie titckeri 
grows on the leaves and fruit of the Vine ; it spins its hyphse over the bunches 
of gtapes, curtails their growth, and causes them to burst, and to become de- 

FIG. 109. Eurotiiim glaucum : a portion of mycelium lying horizontally ; ft vertically- 
placed conidiophore ; the mycelium gives rise to another branch near a ; the conidia are 
abstricted from short flask-shaped cells ; b a ripe conidium ; c, d germinating conidia ; 
e spirally-twisted hypha, commencement of an ascocarp ; / a stage later ; 3 still later, the 
hypha at the base of the coil has given off branches which are applied to it; Ji, i sections of 
young ascocarps. 

cayed and rotten (Grape-disease). The Fungus was first noticed in England 
in 1845, and later was found in all countries where grapes are grown. It is only 
known in the conidial form (" Oidiuni tuckeri"). Many other species of Ery- 
siphe are found on herbaceous plants. Microsphmra has appendages which are 
repeatedly forked at their extremities. M. grossulurice on Eibes grossularia. 



Uncimtla has appendages with spirally-coiled extremities ; on Salix and Acer. 
Phyllactinia has a circle of bristle-like appendages with dilated bases. P. 
guttata on Corylns, Fraxinus, Fayus, etc. 

Order 2. Perisporiaceae, Moulds and Mildews. A group of 
Fungi widely distributed and found in all situations. Usually they 
have a well-developed surface mycelium, and small, round, seldom 
conspicuous ascocarps, containing ovoid, pulley-like spores. They 
are partly saprophytic, partly parasitic, in the latter condition 
having a brown mycelium. 

Eurotium glaucum ( = E. herbariorum, Figs. 109, 110) and E. 
repens live on dead organic matter, preserved fruits, etc. The 
conidial forms of both species are known as " Moulds " (Fig. 109), 
and formerly were described under the name "Aspergillus glaucus." 

FIG. 110. Em-otinm glaucum : a longitudinal section of a half-ripe ascocarp, bounded 
externally by a well-defined layer of cells, enclosing asci in various stages of develop- 
ment; b a semi- ripe, c an almost ripe ascus ; d and e spores seen from the edge and 
tide ; /germinating spore twenty-two hours after been sown in plum juice. 

The conidia for some time remain attached to each other in 
chains (Fig. 109 a) ; they are abstricted from sterigmata arranged 
radially on the spherical, swollen end of the conidiophore. The 
small yellow or brownish ascocarps are frequently found in 
herbaria, especially when the specimens have been insufficiently 
dried. Euaspergillus fumigatus and others are pathogenic, causing 
mycosis in warm-blooded animals. 

Penicillium crustaceum (P. glaucum, Figs. Ill, 112) is an ex- 
ceedingly common "Mould." Its mycelium appears very fre- 
quently on any organic matter which is permitted to remain 
untouched, and soon covers it with a dense mass of blue-green 


conidiophores. These branch at their summits and bear flask- 
shaped cells from which the conidia are abstricted. The ascocarps 
which, both in size and colour, resemble grains of sand, have only 

FIG. 111. Penicillium crustaceum : a conidia (x 300) ; b germination of conidia; c small 
portion of mycelium, produced from a conidium at *, with five conidiophores; d young 
conidiophore ( x 630), a flask-shaped cell is abstricting a conidium; e the same conidio- 
phore after 9-10 hours. 

FIG. 112. Penicillium crustaceum : a two spirally-coiled byphae arise from the my- 
celium, from one of which (archicarp) the asci are produced; b a further step in the 
development of the ascocarp ; the branching archicarp is surrounded by sterile hyphae ; 
c section of young ascocarp ; the larger hyphoe in the centre are the ascogenous 
hyphas ; these are enclosed by a pseudo-parenchyma of sterile hyphas ( x 300) ; d series 
of ripe asci with spores ; e four ascopores seen laterally ; / germinating ascospores 
(x 800). 



been obtained in luxuriant cultivation with a limited supply of 



Capnodium salicinum (Fumago salicina, Cladosporium fumago), a common 
Mildew, forms dark overgrowths on the leaves and branches of various shrubs 
(Poplars, Elms, Willows) and on Hops. The conidia appear in various forms, 
as on conidiophores, in conidiocarps with large multicellular conidia, and in 
couidiocarps with small unicellular couidia ; in nutritive solutions yeast-like 
conidia are also developed. Apiosporium pinophilum produces mildew on the 
leaves of Abies alba and Picea excelsa. (The conidial-forrns were formerly 
described as " Antennaria pinophila "). 

Order 3. Tuberaceae, Truffles. The Fungi belonging to 
this order are entirely subterranean. The mycelium is filamen- 
tous, and partly parasitic upon the roots of plants, especially trees, 
in its neighbourhood ; it is then known as Mycorhiza. The fruit- 
body is relatively large, in some cases about the size of a hen's 

FIG. 113. ruber melanosporum : a fruit-body (nat. size), a portion having been re- 
moved to show the internal structure ; b an ascus with ascospores. 

egg. Internally it is traversed by a number of winding passages 
(Fig. 113 a), the walls of which are coated with the asci. The asci 
(6) contain only a small number of spores, and these are set free 
by the putrefaction of the fruit-body. Conidia are unknown. 

Tuber melanospomm, T. brumale, T. astivum, and other species are edible. 
Terfezia leonis and Charomyces nuvandriformis are also edible. The Truffles 
are always found in woods and under trees, and disappear when these are de- 
stroyed. France and Italy produce the best and the largest number of Truffles, 
which are hunted by specially trained dogs and pigs. 

In Elaphomyces (Stag-Truffle) the fruit-body has a corky external layer, and 
is inedible. Some of the species are found in this country. E. granulatus is 
parasitic on the roots of the Fir. 



Family 3. Pyrenomycetes. 

In this family the hymenium is enclosed in small fruit-bodies, 
peritliecia (Fig. 120 l>), which appear to the naked eye as small dots. 
In shape they resemble a globe or a flask with a narrow mouth, 
through which the spores are ejected (peronocarpic ascocarps). 
Different kinds of reproduction conidia, pycnidia (chiefly with 
microconidia), chlamydospores, and peritliecia are found in the 
same species. The various stages in the life-history of these 
Fungi are so dissimilar, that formally they were considered to be 
different genera. Ergot furnishes a very good example. 

This family may be subdivided into 3 sub-families. 

FIG. 114. A email portion of an ovary attacked 
with Claviceps purpurea (Sphacelia). 

FIG. 115. An ovary with the conidial 
stage of Claviceps purpurea (Sphacelia). 

Sub-Family 1. Hypocreales. 

The peritliecia are pale, flc.shy, brightly coloured, and generally 
aggregated on a stroma. Conidia and chlamydospores occur very 
frequently. Only one order. 

Order. Hypocreacese. In this order the majority are 
parasites upon Flowering-plants (Nectria, Polystigma, Epichloi', 
Claviceps); but some are parasites upon Fungi (Hypomyces, Melano- 
spora), or upon insects (Cordyceps). 

The most important member of this order is the ERGOT (Clavi- 
ceps purpurea, Figs. 114, 115, 116). This Fungus is found in the 
flowers of many species of Grasses, especially the Rye, attacking 
and destroying the ovaries. In the FIRST or CONIDIAL STAGE of the 
attack, the ovaries are found covered with a white, irregularly 



folded mycelium (Fig. 114 m, Fig. 115), formed of numerous 
hyphte woven together and penetrating the wall of the ovary. 
From these a number of liyphse (Fig. 114 a) project into the air 
and abstrict from their apices the conidia (6) which serve as repro- 
ductive organs. The mycelium also secretes a sticky, stinking 
fluid (honey-dew) in which the conidia are embedded in great 
numbers. The honey-dew exudes from the bases of the glumes, 
and is greedily sought by flies, which thus carry the conidia to 
other ovaries. In this manner fresh ears are infected, which might 

FIG. ll&.Claviceps purpurea. A Sclerotium with stromata (cl) (xby 2). V Stroma 
divided longitudinally to show the perithecia (cp). C A perithecium with the surrounding 
hyphee (hy). D An nseus ruptured, with the eight filamentous ascospores emerging. 

escape were the conidia only distributed by the wind. This stage 
formerly was regarded as an independent Fungus, known as Spha- 
celia segetum (Fig. 115). On germination, the conidia produce 
either a new mycelium (Fig. 114 d, c), or new conidia. The SECOND 
or SCLEROTIUM STAGE is the one in which the Fungus passes the 
winter. The mycelium penetrates deeper and deeper into the 
attacked ovaries, their tissues are destroyed and replaced by the 
hypha?, which gradually become more and more felted together. 



A firm, pseudo-parenchymatous mass of hyphoe is thus formed at 
the base of the loosely- woven Sphacelia, which is in part trans- 
formed into the hard sclerotium, and the remainder thrown off. 
A dark, hard, poisonous body, longer than the natural grain, is 
thus formed ; these bodies are known as Ergots, and were formerly 
considered to be a distinct species, Sclerotium clavus (" Secale 
cornutum," Ergot, Fig. 116 A, c). The THIRD STAGE, described 
as Claviceps purpurea, is developed in the following spring from 
the germinating sclerotium, which produces dark-red stromata 
with short stalks. In the stroma numerous perithecia with asci 
and ascospores are produced. The latter may infect young flowers 
of the cer-eals, in which the disease is then developed as before. 

FIG. 117. Nectria cinnabarina : a branch of Acer pscudoplatanus, with conidial-layers 
and perithecia (nat. size); b a conidial -layer (Tuber cularia vulgaris) ; c, a mass of peri- 
thecia. (b and c x 8.) 

Several species of the genus Nectria, with blood-red perithecia, are found as 
dangerous parasites, especially N. ditissima, which causes " Canker " in the 
Beech, Ash, and Apple, etc. ; N. cucnrbitula, which appears on Pine-trees, and 
N. cinnabarina (Fig. 117), whose conidial form was formerly named Tiiberciilaria 
vulgaris. Folystigma rubniin forms shining red spots on the green leaves of 
Prtms-species. -EpicMoe typhina is parasitic on the sheaths of Grasses, on which 
it first forms a white conidial-layer, later on a yellow layer of perithecia. Cor- 
dyccps (Chrysalis Fungus, Figs. 118, 119) lives in and destroys insects, and after 
compassing their death produces the club-formed, generally yellow, stromata, 
one part of which bears conidia (Isaria) and another perithecia. C. militaris 
(Fig. 118) on the chrysalides and caterpillars of moths, is the most common. 



The so-called Botrytis lassiana, which produces the disease known by the name 
of " Muscardine," in silkworms, is probably a conidial form belonging to 

FIG. 118. Cordyceps mi'Htaris. I Stromata with 
conidiophores (Isaria farinosa). II A larva, with 
strornata, bearing per ithecia. Ill A spore. 

'- ! '' fRfc f r I'- 


Fis. 119. Cordyceps robcrtii on tlie larva of Heplalus vlrescens : n stalk of stroma ; b 



Sab-Family 2. Sphaeriales. 

To this sub-family belong the majority of the Pyrenomycetes. 
The perithecia are of a firm consistence (tough, leathery, woody or 
carbonaceous), and of a dark colour. Their covering is quite dis- 
tinct from the stroma when this structure is present. The stromata 
are sometimes very large, and may be either cushion-like, crus- 
taceous, upright and club-like, or branched bodies. In general, 
small, inconspicuous Fungi, living on dead vegetable matter, 
sometimes parasites. Free conidiophores and conidiocarps are 
known in many species, and in several, chlamydospore-like forms 
of reproduction. Orders 3-18 constitute the Spha3riacese of older 

FIG. 120. Striclceriaobducens : a a portion of an Ash-branch with the bark partly thrown 
off; on the wood are numerous black perithecia ( x 20); b longitudinal section through a 
perithecium ; c a spore: d longitudinal section through a pycnidium whose ascospores 
are being ejected ; e portion of the same, with hyphse and spores. 

Order 1. Sordariaceae. Fungi living on dung with fragile perithecia, 
either aerial or buried in the substratum. The dark brown or black spores have 
either a mucilaginous envelope (Sordaria, etc.) or mucilaginous appendages (Podo- 
sjiora), by means of which their expulsion and distribution are promoted. 

Order 2. Chaetomiaceae. Perithecia fragile, free, bearing on the summit a 
tuft of hairs. Chcetomium, on decaying vegetable matter. 

Orders 3-7. Perithecia scattered or aggregated, situated from the commence- 
ment on the surface of the substratum. Stroma wanting. 

Order 3. Trichosphaeriaceae. TricJwsph&ria parasitica (Fig. 121), on 
Abies alba ; Herpotrichia nigra on Picea excelsa and Pinus montana. 
W. B. K 



Order 4. Melanommaceae. Eosellinia quercina lives in the roots of 1-3- 
year-old Oaks, and destroys the plants. 

Order 5. Ceratostomaceae. 

Order 6. Amphisphaeriaceae. Strickeria olducens (Fig. 120) has brick- 
like spores, and lives aggregated on the hard branches of Fraxinus. 

Order 7. Lophiostomaceae. 

Order 8. Cucurbitariaceae. Perithecia tufted, at first embedded, then 
breaking through, often situated upon an indistinct stroma. 

Orders 9-13. The peritheeia remain embedded, and are only liberated by the 
casting off of the covering layers of the substratum. Stroma wanting. 

Order 9. Sphaerellaceae. The species of Spharella have colourless, 
bicellular spores. They live upon the leaves of many plants, and develope 
spherical peritheeia upon the fallen leaves. 

Order 10. Pleosporacese. The conidial-forms of Pleotpnra herbarum and 
P. vulgaris form a black covering on various plants, known as "smuts." 


FIG. 121. TricTiospJwm'a jiarositica : a a twig of Alies alba with epiphytic mycelium ; 
b a leaf with mycelium and sporangia (magnified) ; c a sporangium ( x 60) ; d an ascus 
with spores ( x 650). 

Venturia ditricha occurs on the underside of dry Birch leaves, and perhaps to 
this belongs the conidial-form, Fusicladium pirinum, which causes the "Bust- 
spots " on Apples and Pears. 

Order 11. Massariaceae. 

Order 12. Clypeosphaeriaceae. 

Order 13. Gnomoniaceae. Perithecia, with peak-like aperture. Gnomonia 
erythrostoma in the leaves of Prunus avium, which turn brown and do no fall 
in autumu. 

Orders 14-18. Stroma generally well developed, The peritheeia are embedded 
in the stroma, but ivhen this is rudimentary, in the substratum. 

Order 14. Valsaceae. Valsa. 

Order 15. Diatrypaceae. Diatrype. 

Order 16. Melanconidaceae. 

Order 17. Melogrammataceae. 



Order 18. Xylariaceae. This order is the most highly de- 
veloped of the Sphferiales. The stroma arises on the surface of 
the substratum, which is generally dead or decorticated wood ; 
it is well-developed, crustaceous, hemispherical or upright. In 
the younger conditions it is covered with a layer of conidia, and 
later on it bears the perithecia, arranged in a layer immediately 
beneath its siirface. The ascospores are of a dark colour. Often 
also there are free conidiophores. 

Fia. 122. Xylaria Uypoxyton (nat. size) on a tree stump : a younger, b an older stroma, 
both of which, with the exception of the black lower portion, are covered with white 
conidia ; n, spot where the perithecia are developed ; c an old stroina with upper part 
fallen off ; d, c large branched stromata ; fe conidia. 

Hypoxylon and Ustulina have a cushion-like or crustaceous stroma. Xylaria 
has a club-shaped or branched stroma, often several centimetres high. X. 
hijpoxylon (Fig. 122) and X. polymorplia occur on old tree stumps. Poronia 
grows on old horse dung, and has a conical stroma. 

Sub-Family 3. Dothideales. 

The perithecia are always embedded in a black stroma, and are 
not distinctly separated from it. The accessory forms of reproduc- 
tion are : conidiophores, conidiocarps, and yeast-like conidia. The 
majority are parasites. One order. 

Order Dothideaceae. Phyllachora graminis produces scab-like patches on 
the leaves of the Grasses. Scirrhia rimosa grows on the leaf-sheathes of 
Phragmites. Rhopographus pteridis on Pteridium aquilinum. 



Family 4. Hysteriales. 

This family, like the following, lias hemiangiocarpic ascocarps 
(apothecia). These are closed in the early stages, but when ripe 
open in a valvular manner by a longitudinal fissure ; they are black, 
oblong, and often twisted. Some species are parasites, especially 
upon the Coniferse. 

Order 1. Hysteriaceae. Hysterium pulicare upon the ruptured bark of 
many trees. 

Order 2. Hypodermacese. The species of Lopliodermium live upon the 
leaves of Conifers, and are the cause of their falling off (llifjlit). L. pinastri 


FIG. 123. Lophodenniitm 
(Hypoderma) nervisequium : 
a, two leaves of Abies alba 
seen from above with pyc- 
nidia; b a leaf seen from 
the underside with apothe- 
cia; c an ascus with asco- 
spores. (x 500.) 

FIG. 124. Three leaves of 
the Red-pine with Lopho- 
dermiuin macrosporinn : a 
under side of the leaves with 
apothecia ; b a leaf from 
upper side with pycnidia. 
(x about 2.) 

a b 

FIG. 125. Lopliodcvminm 
pinastri : a leaves of Pinus 
sylvestris with apothecia 
(nat. size);b two paraphyses 
and an ascus with filamen- 
tous spores. 

(Fig. 125), on the leaves of Pinus and Picea ; the leaves become red-brown 
and fall off ; at first conidiocarps are formed, and later apothecia ; L. nervise- 
tlidum (Fig. 123), on Abies alba ; L. macrosporinn (Fig. 124), on Picea e.rcelsa ; 
L. brachysporum, on Pimis strobus. 

Order 3. Dichaenaceae. 

Order 4. Acrospermaceae. 

Family 5. Discomycetes. 

The ascocarps (apothecia) are at first closed, and only open at 
the time of their ripening, not valvularly, but more or less 


like a saucer or cup, so that the hymenium lies exposed on their 
upper surface. In the first three sub-families, and generally also 
in the fourth, the apothecia are formed inside the substratum. 
The apothecia are, in contrast to the Pyrenomycetes, light and 
brightly coloured, and their size varies very much, and may be 
several centimetres in diameter. Paraphyses are often present 
between the asci ; they often contain colouring mattei-, and give to 
the disc its characteristic colour. The tissue on which the asci 
are borne is known as the hypothecium . The shape and colour of 
the spores is not so varied as in the Pyrenomycetes. The acces- 
sory forms of reproduction are conidia (sometimes of two forms), 
chlamydospores, and oidia. The family is divided into 5 sub- 

Sub-Family 1. Phacidiales. 

The apothecia are developed in the interior of the substratum, 
which they break through, and in general dehisce apically. The 
envelope is tough and black. Hypothecium inconspicuous ; 
hymenium flat. 

Order 1. Euphacidiaceae. Phacidium abietimim, on the leaves of Abies 
alba. Rhytisma ; the pycnidia are found in the summer on the green leaves, 
while the apothecia are developed on the fallen leaves and dehisce in the 
following spring. R. acerinum causes black spots on the leaves of the Syca- 
more, and R. salicinum on Willows. 

Order 2. Pseudophacidiaceae. 

Sub-Family 2. Stictidales. 

The apothecia when ripe break through the substratum which 
forms a border round them. Hymenium generally saucer-shaped. 

Order 1. Stictidaceae. Stictis. 
Order 2. Ostropaceae. Ostropa. 

Sub-Family 3. Tryblidiales. 

The apothecia are embedded in the substratum in the early 
stages, and then are raised high above it. Hypothecium thick. 
Hymenium cup-shaped. 

Order 1. Tryblidiaceae. Tryblidium. 

Order 2. Heterosphaeriaceae. Heterospluzria patella on the dead stalks 
of Umbellifers. 



Sub-Family 4. Dermateales. 

The apothecia in the early stages are embedded in the sab- 
stratum and then break through it, or are from the first situated 
on the surface of the substratum. Hypothecium thick. 

Order 1. Cenangiaceae. Cenangium. 

Order 2. Dermateaceae. Dermatea. 

Order 3. Patellariaceae. Patellea, Biatorella, Patellaria. 

Order 4, Caliciaceae. Calicium, Coniocybe, etc., on the bark of trees. 

Order 5. Arthoniaceae. Arthonia on the bark of several trees. Celidium 
stictarum on the apothecia of Sticta pulmonaria. 

Order 6. Bulgariaceas. Apothecia gelatinous tinder moist conditions, and 
horny when dried. Calloria fusarioides ; the red apothecia break out in the 
spring on the dried stalks of Urtica dioica ; a gelatinous reproductive form of 
the Fungus is found before the apothecia, which consists of oidia (formerly 
described as " Dacryomyces urticce "). Bulgaria inquiitans on the living or 
fallen trunks of Oaks and Beeches. 

Sub-Family 5. Pezizales. 

The apothecia are developed on the surface of the substratum and 
are waxy or fleshy ; at the commencement closed, and covered with 
a saucer- or cup-shaped, seldom flat, hymenium. The hypothecium 
is generally well developed. This sub-family is the richest in 

FIG. 126. Botrytis cinerea : a slightly 
magnified ; b more highly magnified ; c 
germinating conidium. 

FIG. 12". Sclerotiniafuckeliania: a scle- 
rotium with conidiophores ; b with apo- 
thecia ; c section through sclerotium and 
apothecium ; d ascus with eight aseo- 
spores. ( x 390.) 



species of the Discomycetes and contains forms of very different 
habit. They grow upon dead wood, upon the ground, and upon 
dung. A few are parasites. 

Order 1. Helotiaceae. Apothecia with waxy envelope of 
colourless, or yellowish prosenchymatous cells. Chlorosplenuim 
ceruginosum is found on decaying wood (particularly Oak and Birch), 
to which it gives a green colour. Sclerotinia has sclerotia which are de- 
veloped upon the host-plant and from which, after a period of rest, the long, 
brown-stalked apothecia arise. S. ciborioides (S. trifoliorum, Fig. 128) is 
parasitic on Clover ; S. sclerotiorum, on Daucus-roots, Phaseolus, etc. ; S. bac- 
carum, on the berries of Vaccinium myrtillus; " Botrytis cinerea " is a common 
parasite and is probably the conidial form of S.fuckeliana (Fig. 127). Helot ium 
herbarum lives on dry plant stems. Dasyscypha willkommii (Fig. 129) produces 
Larch -canker on the bark of the Larch. 



FIG. 128. Sclerotinia ciborioides : a sclerotium 
with three apothecia slightly magnified; b nscus 
with eight ascospores ; c germinating ascospore. 

FIG. 129. Dasyscyphav-illkommii: 
a portion of bark of Larix decidua 
with sessile, cup-shaped apothecia 
(nat. size) ; b two paraphyses on 
either side of an ascus with eight 

Order 2. Mollisiaceae. Mollida cinerea, principally on decaying wood. 

Order 3. Pezizacese. This order contains the largest and 
morphologically the highest forms of the Discomycetes. Apothecia 
fleshy, and in the later conditions generally saucer-shaped. 

Peziza, with sessile apothecia, growing on the ground ; P. cochleata is brown, 
and coiled like a snail-shell ; P. coccinea is scarlet ; P. aurantia occurs as an 
orange-coloured expansion on the ground. 

Order 4. Ascobolaceae. Apothecia fleshy ; in the later stages flat or 



convex. The asci are, comparatively speaking, large, and often contain a great 
number of spores which escape by the casting off of a lid on the summit of the 
ascus. Generally living on dung. Ascubolus furfuraceus, etc. 

Family 6. Helvellales. 

These Fungi have the appearance of clubs, bells, or mushrooms, 
consisting of an upright stalk bearing a large and fleshy head, on 
the exterior surface of which the hymenium is spread. The asco- 

carps are probably gymnocarpic from 
the beginning, and on this account 
these plants are placed in a separate 
family. The development of the 
ascocarps is unknown. The Morchella 
(Morell) grows on the ground ; some 
species are edible. 1 order. 

Order. Helvellaceae. SpathuleaisjeTlovr 

and club-shaped, and forms " fairy rings " in 
woods. Geoylossum (Earth-tongue) projects 
above the ground as a black tongue, or as a 
club-shaped body. Several species are found 
in meadows and on heaths. Helvella has a 
stalk, bearing an irregularly folded head, on 
the external surface of which is the hypothe- 
cium. Morchella (Morell, Fig. 130), the stalk 
bears on its summit the conical or spherical 
head, the external surface of which is reticu- 
late and bears the asci. Mitiula. Verpa. 

FIG. 130. Morchella esculenta: a 
an entire specimen, about one half 
natural size ; b longitudinal section 
through the head. 

Family 7. Ascolichenes (Lichen-forming Ascomycetes). 

The Lichens were formerly classed among the Thallophyta as a 
group quite distinct from the Algae and Fungi. Investigations dur- 
ing the last twenty-five years, however, have conclusively proved 
that the Lichens are Fungi which reproduce in the same manner as 
the Ascomycetes, or, more rarely, the Basidiomycetes, and have 
entered into a peculiar symbiotic relation with Algce, especially 
the Cyanophyceoe and Protococcoideas, with which they associate, 
and without which they would be unable to exist. The Fungus 
forms the largest portion of the Lichen, enclosing the Alga with 
which it may be said to be commensal. The Fungus especially 
produces reproductive bodies and absorbs the inorganic nourish- 
ment through the rhizoids, whilst the Alga supplies it with the 
organic materials. In consequence of this the Lichens, in con- 



tradistinction to other Fungi, need light for the development of 
their nutritive organs, and are therefore, in any case internally, of 
a more or less greenish colour. The form and condition of the 
thallus is unusual among the Fungi, and they can grow upon rocks 
and in other places where no dead organic matter, such as would 
be inquired by other Fungi, is obtainable. 

Two cellular forms are therefore to be found in each Lichen: 
1. The cells which belong to the Fungus. These are generally 
septate, branched hyphfe without any trace of chlorophyll. In 

FIG. 131. Transverse section through the thallus of Skictafuliginosa (x500) : r-r rhizoid- 
strands, which arise from the under side ; g-g gonidial layer ; in medullary layer ; e upper, 
u lower cortex. 

the thallus of the majority of Lichens there may be found a 
medullary layer (Fig. 131 ra) of loosely- woven hyphaa, between 
which there are large air chambers ; and an external layer (cortex) 
(Fig. 131 o, ) formed of closely-woven hyphse without any 
intercellular spaces. In some Lichens (Collemacese) the hypha? 
wind about in the thallus, being equally distributed throughout, 
without forming any decided strata. These Lichens moreover be- 


come gelatinous when exposed to moisture (Fig. 132), on account 
of the swelling of the walls of the Alga?. The hyphas contain 
protoplasm with drops of oil, but never starch ; their walls easily 
swell when exposed to damp after having been dried, and in some 
(e.g. Cetraria islandica) they become gelatinous when cooked. 
Certain strata of hyphas become blue on treatment with iodine 
alone, from which it is inferred that the wall is allied, in its 
chemical nature, to starch. 

2. The enclosed Algae, termed " gonidia." Some belonging to 
the Cyanophycese, Protococcoidese, (especially Pleurococcus) and 

FIG. 132. Collema microphyllum. Transverse section through the thallus ; g Nostoc- 
chains; h hyphse. 

Chroococcacese, are spherical and are found isolated, or in irregular 
groups of cells (Fig. 131 g) ; some belonging to Nostoc (Fig. 
132 </), Lyngbyaceae, etc., are placed in cell-rows. Each Lichen, 
as a rule, has only one definite Algal-form for its gonidium. 

The gonidia either lie together in a certain stratum between 
the cortex and the medullary layer (Fig. 131 g}, or are scattered 
irregularly throughout the entire thallus (Fig. 132). The thallus 
is in the first instance termed " heteromerous," in the second in- 
stance, "homoiomerous." The Fungal-hyphje embrace the gonidia 
and apply themselves closely to, or even penetrate them, and 
hence it has been difficult to decide whether the one cellular form 
does or does not develope from the other (Figs. 134, 135). 



This theory regarding the symbiosis of Fungi and Algae to form a Licheii is 
termed the Schwendenerian theory, after the first scientist who advanced it with 
any weight. It had heen already indicated by De Bary, and further arguments 
in its support have at a later time been adduced by Bornet, Stahl, Treub, 
Frank, Bonnier, Alfr. Holier and others. 

FIG. 131. Nostoc liclienoides, which is attached 
by a germinating thread (?i) of Colletua glaucescens. 

FIG. 133. Ephele pubcfccns. 
The apex of a branch of the 
thallus with two lateral branches 
(*) : h its hypha?;! g the apical 
gonidinrn of the main branch. 

FIG. 135. A Germinating spore of Physcia parie~ 
tina with Protococcus viiidis. B Synalissa eymphorea 
with Glaocupsa. C Cladonia furcata with Proto- 

The thallus of the Lichen appears mainly under three forms : 

1. The CRUSTACEOUS, which adheres firmly to the substratum 

(bark, stone) throughout its entire surface, without being raised 

into any free patches or lobes. It has, in many instances, no 

definite outline, and hyphal-branches from it often penetrate 



deeply into the substratum. It grows at the circumference and 
sometimes dies away in the centre (Figs. 138, 139, 140). 

2. The FOLIACEOUS. This also lies flat upon the substratum, 
but is not firmly attached to and has a definite outline. It grows 

FIG. 136. Portion of a hymenium : A a thin 
stratum on which the asci (s) are situated. 

FIG. 137. Spores of, a Cladonia, Lecanora and Pertusaria ; l> Bceomyccs c Sphinctrina; 
d, e, f various species of Parmelia ; g, h Verrucaria in its younger and older condition ; i, 
fe species of Leptogium. 

at the margin, and raises itself a little by free outgrowths and 
lobes (Fig. 141). The rhizoid-strands spring out from its whitish 
under surface (Fig. 131, r). 

3. The FRUTICOSE, which is attached to its substi-atum at a 
small point from which it projects freely, either erect or pen- 
dulous. It is more or less tufted, in the form of a bush (Figs. 142, 

FIG. 138. Lecanora sitb/i(sca : 
a the bark on which it is situ- 
ated; I the thai I us ; s the 
ascocarp ; s' an ascocarp. 

Fis. 139. Graphis 
(two species). 

FIG. 140. Pertusaria com- 

143). These three thallus-forms gradually pass over by many 
intermediate forms into one another. 

The Lichens, like other Ascornycetes, have very variously 



constructed ascospores (Fig. 137), which are enclosed in asci 
(Fig. 136), usually surrounded by paraphyses attached together. 
Furthermore they possess pycnidia (Fig. 141) containing numerous 
microconidia. These were formerly considered as organs of 
fructification, and were termed " spermatia," and the pycnidia, 
" spermogonia." Alfr. Moller proved, in 1887, that the micro- 
conidia are able to germinate and produce a mycelium with new 
conidia, just as in other Ascomycetes. 

VEGETATIVE REPRODUCTION takes place by soredia, which to the 
naked eye appear as whitish powder on the surface of the thallus. 
They are small round bodies, formed by one or a group of gonidia, 
which are surrounded by a mass of felted hyphas. After the 
rupture of the cortex they are set free, and readily carried by the 
wind to other places, where under favourable circumstances they 

tablish a new thallus. 

FIG. 141. A A portion of the thallus of Parmelia pnrietina with ascocarps (a) and pycnidia 
(I). B A portion of the thallus of Cetraria islandica with pycnidia at the end of small 
lobes. C A lobe with pycnidia and ejected microconidia. (Magnified). 

GEOGRAPHICAL DISTRIBUTION. The Lichens are the most hardy 
plants, and are the first to appear on hitherto bare rocks which 
they gradually disintegrate, and hence prepare the way for the 
growth of other plants. They are to be found from the Polar 
regions to the Equator ; from the highest snow-free mountain- 
peaks down to the level of the sea ; on the stems of trees ; on rocks, 
soil, some even on inundated places ; on stones in woodland streams, 
and on beaches ; but they are never found upon rotten organic 
remains. Some grow gregariously in enormous masses, and form 
wide-stretching carpets, e.g. Reindeer Moss (Cladonia rangiferina), 
species of Cetraria and other fruticose Lichens. 

USES. On account of the cell-wall being composed of Lichen- 


starch (Lichenin), the Iceland-Lichen and Manna-Lichen (Lecanora 
esculenta) are used as food ; the latter grows on stones, in the 
deserts of Asia and North Africa, and is often torn loose in large 
masses and carried away by the wind. The Reindeer-Lichen is 
not only the principal food of the reindeer, but it is also used in 
the manufacture of Danish brandy. Cetraria islandica (Lichen 
islandicus) is OFFICINAL. Colouring materials (lacmus, orseille, 
persio) are made from several species, especially from Roccella 
tinctoria (from the rocky coasts of the Mediterranean). Parmelia 
saxatilis and particularly Lecanora tartarea are used for colouring 
purposes in the Northern countries. 

About 2,000 species of Lichens have been described. If we 
disregard the Basidiolichenes, which will be considered on page 
176, the remaining Lichens (Ascolichenes) may be divided into 
the two following orders according to the structure of the fruit- 
bodies : 

Order 1. Pyrenolichenes. The ascocarps (apothecia) are 
spherical or flask-shaped, as in the Pyrenomycetes, more rarely 
linear (Grapliis) 

According to the nature of the thallus, these Lichens may be divided into : 

a. Thallus homoiomerous, but not gelatinous, branching according to the 
mode of growth of the Algte : Ephcbe (Fig. 133), with Algae of the genus 

b. Thallus homoiomerous, gelatinous : Lichina. 

c. Thallus heteromerous, crustaceous : Verrucaria, Pyrenula; Graphis (Fig. 
139), which may be considered as Hysteriacese with gonidia; several species of 
Graphis are common on bark. 

d. Thallus heteromerous, foliaceous : Endocarpon. 
c. Thallus heteromerous, fruticose : Sphierophorus. 

Order 2. Discolichenes. These, as in the Discomycetes, 
have open apothecia, which, as a rule, are cnpular, more rarely 
hemispherical (Cladonia). 

According to the nature of the thallus, these Lichens may be divided into : 

a. Thallus homoiomerous, but not gelatinous, branching according to the 
mode of growth of the Algae : Ccenogoniiim. 

b. Thallus homoiomerous, gelatinous : Collema (Fig. 132), with Algas of the 
genus Nostoc ; Leptogium. 

c. Thallus heteromerous, crustaceous: Pertusaria (Fig. 140), Lecidca, with 
apothecia open from the beginning ; Lecanora, with apothecia, which in the 
beginning are closed, later on open, but with a rim formed by the thallus 
(Fig. 138) ; Bceomyces, whose apothecia are borne on a stem formed by the 



d. Thallus heteromerous, foliaceous : Parmelia (P. saxatilia P. parietina, 
Wall-Lichen, Fig. 141, is yellow, very frequent on tree-stems, stone-walls, 
tiles); Physcia (P. ciliaris, 
frequent on tree-stems) ; 
Sticta (S. pidmonacea,~Lnng- 
Lichen, on tree-stems) ; -sU^Y 
Peltiaera, especially on the %r^\ rr^N 
Moss among trees ; Umbili- 
caria, on rocks. 

. Thallus heteromerous, 
fruticose : Cetraria (C. ia- 
landica, "Iceland Moss," 
with an olive-brown, flat, 
furrowed, fringed thallus, 
on heaths ; C. nivalis, white, 
in the Polar regions ; 
Evernia, Ramalina, Usnea 
(U. barbata, Beard-Lichen, 
Fig. 143) ; Roccella, Stereo- 
caulon, Cladonia, of which 
the genus C. rangiferina, 
Keindeer-Moss (Fig. 142) is 
important ; Cladonia has two kinds of thallus, one scaly and leaf-like, the other 
erect, which bears the apothecia and maybe fruticose (Fig. 142), or cupular 
(Fig. 144) ; they grow in soil in forests and on heaths. 

FIG. 1-12. Cladonia rangiferina : s ascocarp 

FIG. 143. Usnea barbata : s ascocarp. (Slightly 

FIG. 144. Cladonta pyxidalu. 



Sub-Class 2. Basidiomycetes. 

This sub-class embi'aces the most highly developed Fungi, with 
large " fruit-bodies," which in ordinary language we shortly term 
Funguses, Toadstools, or Mushrooms. 

They have no sporangia, but reproduce only by means of 
basidiospores, conidia, chlamydospores and oidia. The chief 
characteristic of this sub-class is the basidium (Fig. 145), i.e. the 
conidiophore, which has a distinctive form, and bears a definite 
number (generally 4) of characteristically shaped conidia (basidio- 
spores, Fig. 145 c, c/, e). 

FIG. 145. Development of spores in Corticlum. 

The summit of each basidium is produced generally into four 
conical points (sterigmata, Fig. 145 6), from each of which a 
basidiospore is abstricted. The basidia may be classified into 
three principal groups, each of which accompanies a distinctive 
conidiophore: 1, the long, filamentous, transversely divided basidia, 
with lateral sterigmata and spores, found in the Uredinacea3 (Figs. 
146 D, 153), Auriculariacea? (Fig. 160 B}, and Pilacraceae ; 2, the 
spherical, longitudinally divided basidia of the Trernellaceas (Figs. 
160 cd; 161 iii. iv.) ; and 3, the ovoid, or cylindrical, undivided 
basidia of the Autobasidiomycetes (Figs. 145, 163, etc.) ; the two 
last have apical sterigmata and spores. 

The first two groups are the septate basidia (protobasidia), of the Protobasidio- 
ini/cetes; while the unseptate basidia (autobasidia) of the Autobasidiomycetes 
are the third group. On the formation of the basidiospores, the nucleus 


of the basidium divides into four nuclei, each of which ^is transferred to a 

In addition to the basidia, simple conidiophores are also found. 
In the Protobasidiomycetes, the simple conidia are very generally 
found as accessory methods of reproduction in conjunction with 
the basidiospores ; but less frequently in the Autobasidiomycetes, 
e.g. among the Dacryomycetes, Tomentellacege, Heterolasidion 

The simple conidiophores vary in size, and in the number and shape of 
the conidia ; they, however, resemble the basidia, and are doubtless an early 
stage iu the development of the definitely formed basidia. 

Finally, well-defined cldamydospores, formed in various ways, 
appear in the Basidiomycetes as supplementary reproductive bodies 
(compare p. 90). Among the Protobasidiomycetes, chlamydospores 
are at present only found among the Uredinacese, but in various 
forms ; in the majority of families of the Autobasidiomycetes 
oidia frequently occur (Fig. 162), bat genuine chlamydospores 

In the same species several of the known forms of reproduc- 
tion may be distinguished. 

The mycelium is generally composed of white, branched strands, 
consisting of numerous felted hyphas ; in some, sclerotia are 
found. The great majority are saprophytes ; some (particularly 
all the Uredinaceee), are parasites. 


Series 1. PROTOBASIDIOMYCETES : partly gymnocarpic, partly angiocarpic. 

Family 1. DACRYOMYCETES : gymnocarpic. 

2. HYMEXOMYCETES : partly gymnocarpic, partly hemian- 


3. PHALLOIDEJE : hemiangiocarpic. 
4. GASTEROMYCETES : angiocarpic. 
Appended. BASIDIOLICHENES : Lichen-forming basidiomycetes. 

Series I. Protobasidiomycetes. 

To this series belong the lowest of the Basidiomycetes. The 
basidia appear in two principal forms (1 and 2 on page 144) and 
are divided into four cells, either transversely or longitudinally, 
each division forming a sterigma which abstricts a basidiospore. 
The first three orders, Ureclinacete, Auriculariaceae, and Treniel- 

w. B. L 


laceae have gymnocarpic fruit-bodies, while those of the 
Pilacracete, on the contrary, are angiocarpic. 

Order 1. Uredinaceae (Rusts). All the Rust-Fungi are 
parasites, their mycelium living in the interior of the stems and 
leaves of their hosts, causing red, brown, or black spots hence 
their name and malformations, sometimes of considerable size. 

The Rust-Fungi are gymnocarpic and destitute of a hymenium ; 
for these reasons they are regarded as the simplest order of the 
Basidiomycetes. They are entirely parasitic, and their filamentous, 
branched mycelium ramifies in the intercellular spaces of its host, 
and often protrudes haustoria into the cells. The mycelium is 
perennial should it enter a woody tissue ; it may also hibernate in 
the rhizomes of perennial herbs and permeate the shoots springing 
from them, but in the majority of the Rust-Fungi the mycelium 
has a very limited growth. The chief means of reproduction of 
the Rust-Fungi are the chlamydospores, which in the more highly 
developed species occur in three forms, namely, the teleuto-, 
ascidio-, and uredo-spores. The spores, in the host, are formed 
immediately beneath its epidermis, "which is ruptured on the 
ripening of the spores, with the production of " rust," brown, red, 
or black spots. Those chlamydospores which produce basidia are 
termed teleutospores. The spore on germination produces a trans- 
versely divided basidinm, " promycelium," on which basidio- 
spores, " sporidia," generally four in number, are produced on 
lateral sterigmata. This basidio-fructification is gymnocarpic; 
the basidia neither form a hymenium nor a fruit-body (only 
Cronartium and Gymnosporangium have a slight indication of a 

Many Rust-Fungi, in addition to basidiospores, have small, 
unicellular conidia, " spermatia," which are borne in conidio- 
carps, " spermogonia." 

The TELEUTOSPORES ( Winter-spores) maybe either unicellular or 
niulticellular ; in the majority of cases they are enclosed in a hard 
outer cell- wall, the exospore, which in some cases is very strongly 
developed ; they have also a long or short stalk, the remains of 
the spore-bearing hypha. Each cell of the teleutospore has one 
germ-pore (a thin portion of the wall, for the protrusion of the 
germ-tube ; in Phragmidium and Gymnosporangium there are, how- 
ever, several germ-pores). The colour of the teleutospores is 
generally much darker than that of the uredospores, and it is by 
these that the majoi-ity of the Rust-Fungi hibernate. 


In Gymnosporangiurt, two kiuds of teleutospores are found (distinguished 
by their size and thickness of exospore). In many species of Puccinia, the 
form of the teleutospores varies very much, so that in the same layer spores 
have been observed with the characteristic form of other, allied genera. The 
teleutospores of Endophyllnm resemble secidiospores, since they are united 
in chains, whose cells are easily separated, and are produced in the interior of 
a " peridium." The tnulticellular teleutospores of Coleosporium function as 
basidia, and from each cell immediately produce basidiospores. The teleuto- 
spores of Coleospi>rium and Chrysomyxa, differ from other teleutospores in the 
absence of exospore and germ-pore. 

The .ECIDOSPORES (Spring-spores} are produced in chains which 
are generally enclosed in an envelope of hyphae, the peridium ; the 
peridium enclosing the spores being termed the cecidium. The 
secidiospores are unicellular, and generally of an orange colour ; 
they are often separated by intermediate cells which wither and so 
assist in the distribution of the spores. The exospore is made up 
of minute, radially arranged rods. Generally germination proceeds 
immediately, the secidiospore producing a germ-tube, which de- 
velopes into a mycelium bearing either uredo- or teleuto-spores. 

The aecidia of many Rust-Fungi were formerly considered as distinct genera. 
The tecidia of Phragmidium, Triphraymium, and Melampsora, in which the 
peridium is wanting, were in part considered as Cceoma. The ascidia with 
finabriate edge, or those of Gymnosporangium with longitudinal lattice-like 
splits, were considered as " Roestelia " (Lattice-Bust) ; large, sac-shaped tecidia 
on the Coniferae were known as Peridermium. 

The UREDOSPORES (Summer-spores) are unicellular and arise 
singly, seldom in chains (Coleosporiuni). Their colourless, warty 
exospore bears, in the equatorial plane, 2-8 germ-pores. In the 
majority, germination proceeds immediately, and a mycelium is 
produced which at first gives rise to uredospores and afterwards 
to teleutospores. 

The uredospore-formations of Melamp^orella and Cronartium are enclosed in 
an envelope, and hence resemble aecidia. Between the uredospores sterile, 
unicellular hyphae (paraphyses) may be found. 

The spermogonia are spherical or pear-shaped conidiocarps, 
generally embedded in the substratum, and are produced before 
the secidia, before or simultaneously with the uredospores, or before 
the teleutospores. The conidia, as far as observations go, do not 
generally germinate under ordinary conditions. 

Among the Rust-Fungi some species are found which only 
form basidiospores and teleutospores (Puccinia mah-acearum, 


Chrysomyxa abietis). Other species have in addition uredospores ; 
others spermogonia and uredospores; others spermogonia and 
secidia ; others spermogonia, uredospores and aacidia. Those 
species in which all the methods of reproduction are not de- 
veloped mast not be considered as incomplete forms. 

As a rule the mycelium, which is produced from the basidio- 
spores, developes fecidia; in the species, however, without tecidia, 
it developes the uredo-form, and when the uredospores are also ab- 
sent, the teleutospore-forrn. It has been established in some species 
of Puccinia and Uromyces that the formation of fecidia can be sup- 
pressed, and it is not a necessary part of the cycle of develop- 
ment of the species. 

The majority of Bust-Fungi hibernate in the teleutospore-form. Many 
species are able to hibernate in the uredospore-forni (Coleosporium senecionis). 
Others pass the winter in the secidio-form, and develope tecidia on new hosts 
(Uromyces pisi, on Euphorbia cyparissias ; Phragmidium subeorticium, on Rosa ; 
JEcidium elatinum, on Abies alba). In Chrysomyxa abietis, the mycelium, 
developed from the basidiospores, survives the winter. 

Among the Bust-Fungi, with several forms of reproduction, 
there are about sixty whose development can only be completed by 
an alternation of hosts, that is, on one host only uredo- and teleuto- 
spores are produced, while the further development of the ger- 
minating basidiospores, and the formation of the tecidia and 
spermogonia from its mycelium, can only take place on a second 
quite distinct and definite host (lieteroecious or metoxenous Fungi). 
Those Fungi which have all their forms of reproduction on the 
same host are termed autoecious or autoxenous. It is not, however, 
always necessary that the heteroecious Bust-Fungi should regularly 
change their hosts ; for example, Puccinia graminis can hibernate 
in the uredo-form on the wild Grasses, and in the spring can dis- 
tribute itself again in the same form. 

As a consequence of the alternation of hosts the various forms of develop- 
ment were considered as independent genera (Uredo, ^Ecidium, Ramtelia, Ctcoma, 
Peridermium), until De Bary and Oersted established, about the same time 
(1865), the mutual connection of some forms, and paved the way for the right 
conception of these Fungi. 

As an example of one of the most highly developed species, 
Puccinia graminis, the " Bust of Wheat," holds a prominent posi- 
tion. Its uredospores and teleutospores are produced (Fig. 146) 
on Grasses (on cereals, especially Wheat, Bye, Oats, and many 
wild Grasses), while the tecidia and spermogonia are confined to 



the Berberidaceae. The teleutospores, developed on the Grasses, 
hibernate on the dried portions of their host, and in the succeeding 
year each of the two cells of the teleutospore may develope a 
basidium with four basidiospores (Fig. 146 D, c). The basidiospores 
are distributed by the 
wind, germinate quickly, 
and only proceed to fur- 
ther development on Ber- 
beris or Mahonia. The 
germ-tube bores through 
the epidermis of the Bar- 
berry-leaf, and forms a 
mycelium in its interior, 
its presence being indi- 
cated by reddish-yellow 
spots on the leaf. After 
6-10 days the flask-shaped 
spermogonia appear (Fig. 
147 B ; C, a ; conidia in 
Fig. 147 D) and a few 
days later the cup-shaped 
cecidia (Fig. 147 A ; C', c, 
J, e). The former are 
generally on the upper, 
and the latter on the 
under side of the leaf. The 
orange - coloured osciclio- 
spores scatter like dust, 
and germinate only on 
Grasses ; the germination 
takes place in about two 
days when placed on any 
green part of a Grass. 
The germ-tube enters the 
Grass - leaf through a 
stoma ; a mycelium is de- 
veloped in the leaf, giving 
rise to a small, oval, rust-coloured spot (Fig. 146 A) ; in about 6-9 
days the epidermis is ruptured over the red spot, and numerous 
reddish-yellow uredospores, formed on the mycelium, are set free. 
The uredospores (Fig. 146 B) are scattered by the wind, and can 

FIG. 116. Puceinia graminis. 



germinate should they fall on the green portions of other Grasses : 
they then emit 2-4 germ-tubes through the equatorially-placed 
germ-pores. The germ-tubes enter a leaf through a stoma, a 
new mycelium is then developed, and in about eight days a fresh 
production of uredospores takes place, which germinate as before. 
The uredospore-mycelium very soon produces, in addition, the 
brown teleutospores, which give a brown colour to the rust-coloured 
spots, the familiar uredospores on the cereals being quite sup- 
pressed towards the close of the summer (Fig. 146 C, D). The 
" Rust of Wheat " hibernates on some wild Grasses in the uredo- 
s pore- form. 

FIG. 147. JEcidlum lerberidis. A Portion of lower surface of leaf of Barberry, with 
cluster-cups (acidisi). B A small portion of leaf, with spermogonia, from above. C Trans- 
verse section of leaf on the upper side, in the palisade parenchyma are three epermogonia 
(a b); on the lower side an unripe acidium (c d) and two ripe secidia (d, e, /); / chain of 
secidiospores. D Hyphse, forming conidia. 

GENERA. Pitccinia (Fig. 146, 147) has bicellular teleutospores, each having 
a germ-pore, and the aecidia when present have an indented peridium ; some 
species, as exceptions, have 1-3-celled teleutospores. Many species are 
HETEROZCIODS, for example, P. graminis, described above ; P. ntbiyo, which also 
infests various Grasses, but whose tecidia appear on Ancliusa ; the masses of 



FIG. 148. GymnospoTangium, sabince. A 
small portion of the epidermis of a Pear- 
leaf (a) pierced at b by the germinating 
basidiospore (c). 

teleutospores are small ; they contain parapbyses, and are for a long time 

covered by the epidermis. P. coronata, on Oats and Eye Grass ; its tecidia on 

Rhamnus ; the teleutospores are surmounted by a crown "coronate processes." 

P. phragmitis, on Reeds ; ascidia on species of Rumex and Rheum. P. molinice, 

on Molinia ccendea ; the aecidia on 

Orchids. P. poarum, on Meadow- 

Grass ; aecidia on Tussilago. Various 

Pucciuias growing on species of Cares 

have their fecidia on Urtica, Lysi- 

machia, Cirsium, Pedicularis, etc. 

Of those AUTCECIOUS species, which 

have all their generations on the 

same host, may be noted : P. galli, 

P. meuthce, P. viola;, P. epilobii, P. 

asparagi, which grow on the hosts 

from which they have taken their 

specific names. As representative 

of a group which have spermogonia, 

uredo- and teleuto-sporeson the same 

host, but on different individuals, P. 

suaveolens, on the Field-Thistle, 

may be mentioned. The spermogonia have a strong odour. A peculiar group 

(Leptopuccinia) has only teleutospores, which germinate immediately, and 

whilst still attached to their living host. To this group belong P. arenarice, 

on a number of Caryophyllaceaa ; and P. malvacearum, on various Malvaceae, 

introduced iu 1873 from South America to Europe, where it soon proved very 

destructive to Hollyhocks. 

Uromyces (Fig. 149) differs only from Puccinia in always having unicellular 
teleutospores. Among this genus both hetercecious and autcecious species are 
found. To the first group belong U. pisi, whose aacidia are found on Euphorbia 
cyparissias, and U. dactylidis, whose aecidia appear on Ranunculus ; to the 
second group belong U. beta, U. pliaseoli, U. tiifolii. 

Triphraginium has teleutospores with three cells (one below and two above), 
on Spircea ulmaria. 

Phragmidium (Fig. 150) has teleutospores consisting of a row of cells (3-10) 
arranged in a straight line ; the upper cell has one 
germ-pore and the others four germ-pores placed 
equatorially. Both this and the preceding genus 
have large, irregular aacidia without peridia, but often 
with bent, club-like paraphyses (150 b and c) ; they 
are all autcecious. and are only found on the Eosaceaa. 

Endophyllum (see above, under teleutospores, p. 
147) on species of Semperrivum. 

Gymnosporangium (Figs. 152, 154) has bicellular 
teleutospores collected in large, gelatinous masses 
formed by the swelling of the long spore-stalks ; in each cell 2-4 germ-pores are 
found. Uredospores are wanting. All the species are hetercecious ; the teleuto- 
spores appear on Junipcrus, the aacidia (Rccstelia) on the Pomaceae. G. sabince, 

FIG. 149. Uromyces 
genista ; a uredospore ; b 



on Juniperus sabina, ,T. virginiaua, etc., has the secidia (" Rastelia cancellata ") 
on Pints communis (Figs. 152, 148) ; G. juniperinum, on Juniperus communis 
with " Ecestelia cornuta" (Fig. 154 a) on tiorbus auciiparia, Aria nivea (S.aria) 
and Mains communis; G. clavariesforme on Juniperus communis, the secidiura 
belonging to it (" Emstelia laceruta ") on Crategus oxyacantha. 

Melampsora has prismatic teleutospores placed parallel to each other and 
forming a crustaceous layer ; in many species they are divided longitudinally 
into several cells (Fig. 151). The ascidia, without peridium, belonged to the old 
genus Ceeoma. M. caprearum, on Willows, has the fecidia (Ctcoma euonymi) on 
Euonymus. M. hartigii, on Osiers ; the ascidiurn on Eibes. M. mixta, on Salix 

FIG. 150. Phragmidivm gracile : a an uredospore ; 6 and c two paraphyses; da young 
teleutospore ; e a teleutospore with a basidium and two basidiospores (s) ; / two series of 
secidiospores (Ph. rosce). 

repens and Orchids. H. pinitorqua, on leaves of the Aspen, recidia on Pine 
branches (Pine shoot fungus) ; M. populina on Po/n/Z.s monilifera and nigra ; 
M. letulina (Fig. 153), on Birch leaves ; M. padi (Fig. 151), on leaves of Pnmus 
padus, developes teleutospores in the epidermal cells ; M. lini is the cause of 
injury to the Flax ; M. agrimonue. 

Calyptospora gaeppertiana ; teleutospores on Vaccinium vitis idcea ; spermo- 
gonia and aecidia on Abies alba (Firneedle-Eust). 

Coleosporium (Fig. 155) forms its m-edospores in reddish -yellow chains; for 



the teleutospores, see page 147. C. senecionis, on the Groundsel ; its aeculium 
(Peridermhtm ivolffii] on Pine-leaves (Fig. 155 a). Other species on SoncJnis, 
Petasites, Campanula, Jthinantliaccce. 

Chrysomyxa (Fig. 156) has bright red, branched teleutospore-chains ; each 
spore developes a 4-celled basidium. C. ledi, on Ledum palttstre ; its fecidia 
on the leaves of the Fir. C. alietis (Fig. 156), without uredo- and secidio- 
spores ; teleutospores on the leaves of the Fir. In the first summer, yellow 

FIG. 151. MeJampsora padl: a andb 
uredospores ; c-f teleutospores, seen 
from different sides. 

Fig. 152. Pear-leaf, seen from the under 
side, Kilh"R(esteliacancellata": indifferent 
ages (a youngest, d oldest). 

FIG. 153. Melampsora lietulina : a 
uredospores ; b three contiguous teleu- 
tospores, one of which has developed 
a basidium with three basidiospores. 
(x 400.) 

bands are formed on the leaves, and in the following spring the red cushions of 

Cronartium (Figs. 157, 159) has unicellular teleutospores united in numbers 
to form erect threads or columns; the uredospores are enclosed in a "peridium" ; 
C. ribicola (Fig. 157), on leaves of Eibes (especially Black Currants) ; its tecidia 
(Peridermium strobi, or P. klcbahni) on the stems and branches of Pinus strobus 



FIG. 154. Gymnosporanginum juniperinum : a a smal leaf with three clusters of fecidia 
(nat. size) ; b three conidia ; c two fecidiospores on one of which are seen the germ-pores ; 
d a portion of the wall of an secidium ; e, f two teleutosporee. 

FIG. 155. Coleosporlum senecionis ; a Pine-leaves with fecidia (Peridermium wolffii) nat. 
size; b an secidiospore ; c a germinating aecidiospore ; d a chain of uredospores : e a chain 
of teleutospores of which the terminal one has germinated and produced a basidiospore (). 



(Fig. 159), on which it causes great damage ; C. asclepiadeum, on Vincetoxicum 
officinale; its fecidia (Peridermium cornui) on the stems and branches of Pinus 


FIG. 156. Chrysomyxadbieti*: 
a leaf of the Fir, with 5 clusters 
of basidiospores( x 4); b branched 
rows of teleutospores springing 
from the mycelium (m). 

FIG. 157. Cronartium rlbicola : a mass of uredo- 
spores ( x 50) ; b an uredospore ; c a column of teleu- 
tospores (x 60); da small portion of the same 
more highly magnified, with a basidium and two 
basidiospores (s). 

To the Fungi of which the aecidium is known, whilst the remaining forms are 
still undetermined, but which are without doubt hetercecious, belong -Ecidium 
elatinum, which produces the enormous " witches' brooms " and barrel-shaped 
swellings on stems and branches of Abies alba; and JEcidium strobilinum 
(Fig. 158), which attacks Fir-cones, 
causing all the scales to become 
covered with clusters of aecidia 
opening by a lid. Hemileia vasta- 
trix destroyed the coffee planta- 
tions in Asia. 

Order2. Auriculariaceae. 

The long, transversely divided 
basidia bear laterally 4 long 
sterigmata with basidiospores 
(Fig. 160 J5) and are united 
to form an hymenium on the 
surface of the fruit -body. 
Parasites or saprophytes. 

FIG. 158. -iEcfduim strobilinum : a scale of 
cone of Picea excelsa, with numerous aecidia ; b 
tecidiospores arranged in a series; c a cell 
of the peridium. 



Auricularia sanibuclna (Auricula judte), Judas'-ear, has large fruit-bodies, 
which may attain the size of several inches, resembling an ear or a mussel shell. 
In the moist condition they are flesh-coloured, tough and gelatinous, but when 
dried, become hard, grey and wrinkled ; the exterior is covered with short hairs ; 
while the internal surface bears the hymenium. Habitat : stems and branches 
of old Elder-trees (Sambucus). 

Order 3. Tremellacese. The round, pear-shaped, longitudinally 
divided basidia bear 4 elongated sterigmata, situated apieally, and 4 

basidiospores (Fig. 160 C, I)}, and are united 
into the hymenium on the surface of the 
fruit-body. The fruit-bodies are frequently 
gelatinous and quivering ; similar fruit- 
bodies are also found in the Dacryomyce- 
tacete and Hydnaceaa. Simple conicliophores, 
which appear not nnfrequently in the basi- 
diocarps, before the basidia, are known in 
many species. Saprophytes. 

Exidia has kidney-shaped, oblong basidiospores, 
and small, hook-like conidia ; E. ylandulosa, E. albida, 
etc., on wood. Craterocolla has couidiocarps ; C. cerasi 
on Cherry-wood. Sebacina iiicrustans ; the yellow, 

FIG. 159. Peridermium 
strobi : a?cidia of Cronar- 
tium, ribicola (nat. size). 


FIG. WO.SAv.rieularidstimbueina: a-d basidia in vari- 
ous stages of development; e a sterigma bearing a spore. 
C Tremella lutescens : a-d basidia seen from various sides 
(b from above) and in various stages of development; 
e sterigma with basidiospore (x 400). D Exidia glandu- 
losa : a-c various stages in the development of a basi- 
dium ; d sterigma with basidiospore (x 350). 

fleshy, or cartilaginous fruit-bodies are found in autumn covering the ground in 
moist woods. Tremella has round basidiospores ; T.mesenterica has irregularly- 



folded, quivering, orange fruit-bodies, about one inch in breadth ; T. lutescens 
(Fig. 161) has orange-yellow conidial- and yellow basidial-layers ; T. frondosa 
has fruit-bodies upwards of a foot in breadth. 

Order 4. Pilacraceae. The transversely divided basidia have no 
sterigmata, but sessile basidiospores, and fill up the cavity of a closed 
(angiocarpic) fruit-body as a gleba without a regular arrangement 
(hymenium wanting). 

Pilacre fagi on the old stems of the Copper-Beech ; P. pctersii, on dried 
branches of the Hornbeam, has stalked, capitate fruit-bodies. 

- ---< . 

.'/r t - J ' ." ^ -,5--- 'W' 


On "'' T 23T ' '^ x ri 

FIG. 161. Tremella Itit?scfits: I and II fruit-bodies (nat. size); III vertical section 
through a fruit-body ; b basidia ; c conidia; IV- VI basidia; VII basidiospore with a 
second spore ; VIII a basidiospore with yeast-like budding (cultivated) ; IX a conidio- 
phore. (III-IX about 400.) 

Series 2. Autobasidiomycetes. 

This second and larger part of the Basidiomycetes is character- 
ised by its more highly differentiated, undivided, club-shaped, or 
cylindrical basidia, which generally bear 4 (seldom 2, 6, 8) apically- 
placed sterigmata and basidiospores (Fig. 145). The fruit-bodies 
are partly gymnocarpic (in the first 3 orders and in some Agarica- 
cece). partly hemiangiocarpic (in orders 3-6 of the Hymenomycetes 



and in the Phalloideas, the fruit -bodies in these orders are in the 
young conditions more or less angiocarpic, but later on generally 
open below and bear the hymenium on the under surface of the 
fruit-body), partly also angiocarpic (in the Grasteromycetes). 

FIG. 162. Dacryomyccs deliquescens : I fruit-body (nat. size); II vertical section through, 
the hymenium; III germinating basidiospore ; IV a portion of mycelium with conidia ; 
V a germinating coiiidium ; VI and VII chains of oidia more or less strongly magnified ; 
VIII basidiospore of D. longisporug ; IX germinating basidiospore of D. ovisporus ; X 
and XI Calocera vifcoxa; X fruit-body (nat. size); XI basidia with basidiospores (highly 
magnified); XII Dacryomitra glossoides (nat. size). 


Family 1. Dacryomycetes. 

The long, club-shaped basidia bear two tape-ring sterigmata, which 
develope remarkably large basidiospores (Fig. 162 II, XI) and 
form gymiiocarpic fruit-bodies with hymenium. 1 order : 

Order 1. Dacryomycetacese. This order comprises 4 genera 
of which the first two develope the hymenium on the whole surface 
of the fruit-body, but the two last only on its apex. 

Dacryomyces : the folded, gelatinous, Tremella-\ik& fruit-bodies break out in 
winter on dried wood (hedges) in the form of red or yellow drops. D. deliques- 
cens is very common (Fig. 121). The following genera have cartilaginous 
fruit-bodies. Calocera (Fig. 162), with club-like, simple, or branched, Clavaria- 
like, fruit-bodies ; the orange coloured fruit-bodies of C. viscosa grow aggregated 
together on the wood of Conifers. Guepinia resembles a Peziza, and has the 
hymenium only on the hollow upper surface. Dacryomitra resembles a Hit- 
rula (Fig. 162). 

Family 2. Hymenomycetes. 

This family is very rich in species (more than 8000 have been 
described), and to it belong all the " Mushrooms " and " Toadstools." 
The fruit-bodies present very various forms ; they are generally 
fleshy, very perishable, seldom leathery or corky, in the last case 
often perennial. The basidia are more or less cylindrical and bear 
generally 4 (seldom 2, 6 or 8) sterigmata and basidiospores. The 
hymenium in the fully-formed fruit-bodies lies free on the sur- 
face : in orders 1 and 2 and a portion of order 6 it is from the 
commencement exposed, fruit-bodies gymnocarpic ; orders 3-6 have 
hemiangiocarpic fruit-bodies (p. 157). In the first order the 
basidia (or the hymenium) are developed immediately from the 
mycelium (Fig. 163); the fruit-bodies of orders 2 and 3 present a 
higher grade of development, and have between the mycelium and 
hymenium a special hyphal-tissue, a stroma, which is crustaceous, 
club-like, or coralloid, etc., and in general bears the hymenium on 
the largest part of the free, smooth surface. In the forms most 
highly developed (orders 4-6) a new tissue the hymenophore is 
introduced between the stroma and hymenium, which appears on 
the under side of the fruit-body in the form of warts, projections, 
tubes, folds or lamellae (Figs. 166, 167, 174 be). Paraphyses are 
frequently found in the hymenium, among the basidia. In the 
Hymenomycetes few examples of conidia can be recognised at 
first. More fi-equently chlatnydospores are found, particularly oidia. 
The mycelium is richly branched, generally colourless, often peren- 
nial ; it lives in humus or decaying wood, and is seldom parasitic. 



The hyphre generally have clamp-connections and unite, some- 
times, to form a rhizomorpha (Fig. 177) or sclerotia with coloured, 
pseudopai-enchymatous covering. 

FIG. 1G3. Exobasidium vaccinii. I Hypertrophied stem of Vaccinium vitis idcea ; II lea 
with gall-like swelling ; III section of II ; IV transverse section : m mycelium between 
tlie parenchymatous cells ; p hypodermal cells ; e epidermis with basidia in various stages 
of development ; V epidermis with germinating spores; VI and VII spores germinating 
in water (IV-VII x 620). 



Order 1. Tomentellaceae. To this order belong the simplest of 
the Hymenomycetes. The basidia (Fig. 145) arise free and irregu- 
larly from the mycelium ; a Injmenium is entirely absent or very 
slightly formed (in Corticium it attains its highest development); 
fruit-bodies are also wanting. -In general they form flaky, mem- 
branous or leathery coverings on bark and wood. Some 


Hypoehnus without conidia. Tomentella with conidiophores ; growing on 
wood or earth. Exobasidium vaccinii (Fig. 163), a parasite on Vaccinium, 
Andromeda, Arctostaphylos, and Rhododendron, forms flaky-powdery, white or 
red coverings and may cause hypertrophy of the parts attacked. E. warmingii 
is parasitic on Saxifraga E. lauri causes outgrowths on the stem of Laurus 
canariensis as long as a finger, which formerly were regarded as aerial roots. 
Corticium forms membranous to leathery layers or crusts ; C. quercinum on 
wood and bark, particularly Oak, is flesh-coloured ; C. ccenileum has a blue 
hyrneniurn ; C. giganteum on the bark of fallen Pine-trees. 

Order 2. Clavariacese. The hymenium is situated on a stroma, 
and either completely covers the smooth surface of the more or less 
fleshy gi/mnocarpic fruit-body, 

or is confined to a tolerably VM vu 

well defined upper portion 
of it (Typhula). Paraphyses 
absent. The vertical, white, 
yellow, or red fruit-bodies 
are roundish or club-like, 
undivided or richly branched 
(Fig. 125). Generally on the 
ground in woods, seldom on 
tree-stems, etc. 

GENERA : Clavaria, generally 
large Fungi with thick, round 
branches. C. botrytis has a very 
thick, tubercular stem with numer- 
ous short, flesh-coloured branches: 
it has an agreeable taste. C. coral- 

lo'ules has a brittle, richly-branched fruit-body (Fig. 164) ; basidia with two 
large spores. C. pist illaris consists of a single, undivided club of a yellowish- 
white colour. Sparassis has compressed, leaf-like, curled branches ; S. crispa 
has fruit-bodies as large as a white cabbage-head, with an agreeable taste. 
Typhula and Pistillaria are small Fungi with filamentous stalks, terminating 
in a small club. The fruit-bodies of the former often arise from a small, 
spheroid sclerotium ; the latter is distinguished by the basidia bearing only 
two spores. 

W. B. M 

FIG. 161. Clavaria coralloides (nat. size . 


Order 3. Thelephoraceae. The hymenium is placed on a 
stroraa and covers the smooth surface of the leathery hemiangio- 
carpic fruit-body, generally on its under side. The edge of the 
stroma, which bounds the hymenium is sometimes especially 
developed (Stereum). Saprophytes. 

GENERA: Thelephora. The fruit-bodies in this genus are brown, very ir- 
regularly shaped, and often lobed. The spores too are brown, but in the other 
genera colourless. The species are found growing on barren soil. T. 
laciniata (Fig. 165) has imbricate, semi-circular, dark-brown pileus, which is 
jagged at the edge and upper surface. The fruit-bodies are very often raised 
above the grund, and although this species is not a parasite, yet it destroys 
young seedlings by growing above and smothering them. Stereum has a 
stiffer fruit-body, with a distinct, fibrous, intermediate layer. It grows on 
bark and wood, projecting like a series of imbricate brackets. S. hirsittum is 
yellow ; its free edge is provided with a number of stiff hairs, the upper surface 
being divided into a number of zones. S. purpnreum has a red-violet hymenium 

which distinguishes it from the 
previous species. Cyphella has a 
membranous cup, or bell-shaped 
fruit-body, often borne on a stalk, 
the concave surface being covered 
with the hymenium. They are 
small, white Fungi, growing on 
Moss and dead stems. Solenia is 
closely related to Cyphella ; its 
fruit-bodies are smaller and hairy ; 
they are found clustered together 
forming a crust-like covering on 

a, ,~. TT. T i dead wood. Craterellus has a 

FIG. 16o. Thelephora ln.cvma.ta (nat. size). 

large, funnel-shaped fruit-body, 

the hymenium covering the external surface. C. cornucopioides is shaped like 
a trumpet or a " horn of plenty." It is dark-grey, several inches in height, and 
grows gregariously on the ground in forests. It is distinguished by the basidia 
bearing only two sterigmata. 

Order 4. Hydnaceae. The fruit-body is most frequently 
fleshy, and varies considerably in shape, the simplest forms being 
resupinate, 1 the higher ones umbrella-like. The hymenophore is 
found on the free or downward-turned surface, and always takes 
the form of soft emergencies hanging vertically downwards. The 
emergencies may be thorn-, awl-, or wart-like. The species are 
found growing on the soil and on dead wood. 

GENERA : Hydnum has subulate, distinct emergences. H. repandum is 
yellow, the stalk being placed in the centre of the pileus. It is an edible 

1 In the resupinate fruit-bodies a fertile and sterile surface cannot be dis- 
tinguished (cf. Polyporacese and some Stereum-species). 



species, and 'often forms " fairy rings " in woods. H. auriscalpium (Fig. 166) 
is dark-brown, with stalk placed at the edge of the pileus. It grows on old Fir- 
cones. H. erinaceus grows on old tree-trunks. The fruit-body is yellow and 
very large as big as a human head with emergences as much as an inch in 
length. Irpex has a leathery fruit-body, partly resupinate, partly with free, 
projecting edge ; the under side bears tooth-like emergences which are arranged 
in rows, and Irpex thus forms a transition to the Agaricacese. Phlebia is entirely 
resupiuate, with radially-arranged folds on the free side, and pectinate border. 

FIG. 106. Hydnum auriscalpium, upon a Fir-cone, in different stages of development. 

Order 5. Polyporaceae (Pore-Fungi). An order very rich 
in species (about 2000 species are described). The fruit-body is 
of very different forms resupinate, projecting like a bracket, 
hoof-like, or umbrella-shaped. In some it is fleshy and edible, 
in others leathery or corky, persisting for several years. The 
hymenophore is situated on the under side of the fruit-body, and 
consists of wide or narrow tubes or pores, whose inner surface is 
clothed with the hymenium (Fig. 167). In some fruit-bodies large 
cavities are to be found, which have arisen as interstices between 
the labyrinthine curved and reticulate folds. Chlamydospores are 
known in some species. Conidia occur very rarely. Many species 
work considerable damage : some as parasites on trees, others by 
destroying timber. 

GENERA. Poly poms (Pore-Fungus). The tubes are narrow, accurately fitted 
together, and forming a thick layer on the under side of the fruit-body, appearing 



as a number of fine holes. The fruit-body most frequently resembles a bracket, 
or is hoof-shaped, with one side growing from a tree-trunk ; it is very often per- 
ennial, and a new layer of tubes arises in each succeeding period of vegetation. 
Strata, corresponding to the periodically interrupted growth, are thus formed in 
stories one above the other, and are visible on the upper surface of the fruit- 
body, as well as in the interior, as a series of concentric belts, sometimes as 
many as half a score or more in number. P. fomentarius (Touchwood) attacks 
trees, especially the Beech. The spores germinate on wounds from broken 
branches, and the hyphse, following the course of the medullary rays, find their 
way into the interior of the tree, from whence the mycelium spreads upwards, 
downwards, and peripherally, so that the wood becomes rotten (" white-rot ") 
and thick felts of mycelium are formed in radial and tangential directions. A 

dark line, caused by the 
youngest parts of the hy- 
phae containing a brown 
juice, marks the boundary 
between the rotten and 
the unattacked parts of 
the stem (Fig. 168); at 
places where the mycelium 
extends to the bark, the 
cambium becomes de- 
stroyed and further growth 
is arrested, so that longi- 
tudinal furrows arise on 
the stem. It is at these 
places, too, that the Loof- 
shaped, ash - coloured 
fruit-bodies are developed, 
which may attain a cir- 
cumference of upwards of 
7 feet. The interior of 
the fruit-body consists of 
a dried-up, loosely felted, 
red-brown mass of hyphffl, 
which has been used for 
tinder and as a styptic 
("Fungus chirurgorum"). 
P. igniarius has a harder, 
dark-brown, more rounded 
fruit- body ; it grows in a 
similar manner, but 

especially attacks Oaks, Poplars, and Plum-trees, the wood of which becomes 
rotten, and is called touchwood. P. pini (Trametes pint), (Fig. 170), a parasite 
on the stems of Pinus, causes a kind of " red-rot " in the stem. P. sulphureus 
has a soft, cheesy, yellow fruit-body; it produces " rot" in Oaks and Apple- 
trees. P. offlcinalis, Larch-fungus (" Fungus Laricis " in Pharmocopo3ia), grows 
on Larch-trees in the south-east of Europe. P. versicolor has thin, semicircular 

FIG. 167. Polyporus igniarius. Section through the 
under side of the Fungus : 7i-ifi is hyphal-tissue between 
the tubes, formed by irregularly felted hyphfe, many of 
which are seen cut across; s is the hymenium which 
covers the walls of the tabes, and from which the basidia 
with the spores protrude. 



fruit-bodies, with zones of various colours on the upper side ; it is one of the 
most frequent species on tree-stems. P. frondosus grows on soil in woods, and 
consists of numerous aggregated fruit-bodies, which become very large and 
fleshy. This species is edible. P. perennis also grows on the soil in woods ; 
it is very leathery, with central stalk, and has concentric zones on the upper 
surface of the fruit-body. P. vaporarius destroys the wood of living Pines 
(Pinus silvestris) and 
Firs (Picea excelsa), 
causing it to become 
red-brown ; in timber 
this Fungus causes 
"red-strip" followed 

^^-V-^^AX^^.,^-^-^ "** j 

by a "dry-rot." P. 
squamosus destroys 
many Walnut - trees, 
and is also very de- 
structive to Limes and 
Elms. P. fulvus causes 
a "white-rot" in Abies 

Heterobasidion an- 
nosum (Polyporus an- 
nosus, Trametes radici- 
perda, Fig. 169) is 
characterized by its 
Aspergillus-like coni- 
diophores. It is a 
parasite on the Pine, 
Fir, Birch, Beech, etc., 

FIG. 168. Section of stem of a Beech attacked by P. /omen- 
tarius : a non-attacked parts of the stem ; b the furrows where 
the mycelium has reached the bark, and where the thick 
mycelium-strands reach the exterior (J-th of the nat. size). 

FIG. 169. Base of a Fir-tree, with a number 
of fruit-bodies of Heterobaaidion annosum just 
beneath the surface of the soil, indicated by 
the dotted line (jth nat. size). 

FIG. 170. A fully developed fruit- 
body of Polyporus jnni (Trametes 
vini), lateral view (nat. size). 



and is the chief cause of a root-disease (red-rot) in Pines and Firs ; the fruit- 
bodies develope a large number of basidiospores ; they may be very large and 
are found just beneath the surface of the soil (on living or dead roots), and 
exposed to the air (on felled stems and roots, in Scandinavia). 

Ptychogastcr has cushion-like fruit-bodies, which consist chiefly of chlamy- 
dospore-chains, formed of ellipsoidal spores, which alternate with short hyphse 
having transverse septa and clamp-connections. The hymenial portion is 
limited to a small group of tubes. Ft. albus (Oligorus ustilaginoides) grows on 
stumps of Conifers and forms irregular cushions, at first white and later on 
brown, which consist almost entirely of chlamydospores. 

Boletus (Fig. 171) has a fleshy fruit-body resembling a common Mushroom, 
with central stalk. The layer of tubes is easily detached from the pileus, and 
the tubes are easily separable from one another. They grow on the ground in 
woods. Edible species are: B. edulis, with thick, reticulate stalk; B. scaber, 

with thin stalk and rough pileus ; B. luteus, with 
a ring on the stalk. B. luridus is poisonous, 
its tubes have red openings, and the flesh turns 
quickly blue when broken and exposed to the air. 
Fistulina hepatica (Beef-steak Fungus), has a 
red, fleshy, edible fruit-body, with red juice. 
The tubes are individually distinct ; conidia are 
also developed. Grows on old Oaks. 

Merulius lacrymans (" Dry-rot") has a resu- 
pinate fruit-body with white, cotton-like border, 
and the remaining portions covered by reticulate, 
ramified veins of a rust-brown colour. In favour- 
able vegetative conditions it is fleshy and exudes 
large drops of water hence its specific name 
and also the name " Tear Fungus." The my- 
celium is at first colourless, and then yellow- 
brown ; when dry it is tough and leathery. It 
destroys the timber in damp houses, extends far 
and wide over boards and beams and even over 
the masonry, giving rise to a disagreeable smell 
in the rooms in which it lodges. In woods the 
Fungus lives on Pine-stems. It is brought from the forest on the logs of timber, 
and is distributed from log to log by the mycelium and the basidiospores. 
The living mycelium can be recognised by the clamp-connections shooting out 
branches. The basidiospores are often ejected a distance of a metre ; they are 
elliptical (10-11 /j. long and 5-6 fi broad), and germinate easily on damp wood, 
or in fruit-juice which has been neutralized with urine or alkaline carbonates. 

Dtfdalea (Labyrinth Fungus), has bracket-like, corky fruit-bodies with irregu- 
larly-folded plates or discs on the under side. It forms a transition to the 
Agaricaces. D. quercina is frequent on Oak- stumps. 

Order 6. Agaricacese (Mushrooms, Toadstools). The 

hymenophore consists of knife-like plates (lamellae, gills), which are 
situated on the under side of the umbrella-like pileus of the frnit- 


FIG. 171. Boletus edulis (about 
Jth) : b, longitudinal section of 
a portion of the pileus. 



body, and radiate from the central stalk. Those which are first 
formed extend from the edge of the pileus to the stalk; those 
formed later reach only a longer or shorter portion of this distance, 
according to their age. In structure the lamellae (Fig. 174) con- 
sist of a central mass of hyphas, the trama, continuous with the 
hyphae of the pileus ; these terminate in a layer of shorter cells, 
the subtly menial layer, immediately beneath the hymenium which 
is composed of basidia and paraphyses. In a few species, but not 
in the majority, the lamellae are branched, and in some they are 
decurrent. A few have the stalk placed excentrically, or it may 
be entirely absent. 

FIG. 172. Development of Psalliota campestris : a, b, c, d show the various stages of 
the development of the fruit-bodies and the mycelium (in) (nat. size); the fruit-body 
in a somewhat later stage, slightly magnified ; / longitudinal section of e ; n first forma- 
tion of the hymenium ; g longitudinal section of a more advanced fruit-body (nat. size) ; 
n the hymenium ; o velum partiale (see Fig. 133.) 

In the early stages of its development the fruit-body is more or 
less enclosed in a hyphal tissue the " veil " (velum universale, or 
volva). The veil at first completely encloses the young fruit-body, 
but is afterwards ruptured as the latter grows, part remaining 
at the base of the stalk as the "sheath" (annulus inferiis), and 
part on the pileus as scales or warts. In the " Fly Mushroom " 
(Amanita muscaria) the remains of the veil are especially con- 



spicuous as white patches on the bright red ground of the upper 
surface of the pileus, and as a sheath at the base of the stalk (Fig. 
178 r.). Another veil the velum partiale -a hyphal tissue (Figs. 
178 a; 173) stretches from the edge of the pileus to the stalk, and 
encloses the lamellae. This veil is ruptured as the pileus expands, 
a portion attached to the stalk remaining as the " upper ring " 
(annulus superus) (Figs. 173, 178 a), or a part attached to the 
pileus hanging down as a fringe round its edge. Some genera 
have no veil, the under side of the pileus being exposed from the 
first (yymnoc.arpic Agaricacea?). Those which have a veil (hemi- 
angiocarpicA.) afford a transition to the angiocarpic Grasteromycetes. 

FIG. 173. The cultivated Mushroom (Psalliota campe&tris). 

The mycelium mostly grows in soils rich in humus or dung, on 
decaying trees and similar objects. Many species, e.g. Triclioloma 
personatum and Marasmius oreadcs, form the so-called " fairy rings." 
The fruit-bodies in these species are confined to a larger or smaller 
surface on which they are very regularly arranged in a ring. The 
reason for this is found in the radial growth of the mycelium, so 
that the oldest portion, or the starting point, is found at the centre 
of the ring, and the younger ones, on which the fruit-bodies are 
formed, at the circumference. The older hypha? gradually die, and 
at the same time, the radial growth continuing, the ring of fruit- 
bodies becomes larger and larger. The " fairy -rings " are marked 



not only by the fruit-bodies, but also by the more vigorous growth 
and darker colour of the grass upon these spots. 

Some species are parasites. An example is presented by Armil- 
laria mellea, a remarkable and very destructive Fungus in woods and 
forests (Figs. 176, 177). In addition to the filamentous, white mycelium, it 
has also black, or black-brown, horny, root-like mycelium-strands (rhizomorpha) 
which were formerly 


considered to belong to 
a special genus of Fungi 
described under the 
name " Rhizomorpha." 
The mycelium lives 
parasitically on the Con- 
ifers and other trees, 
forcing its hyphas into 
the bark and between 
the bark and wood, and 
thence penetrating into 
the wood so that the 
tree is very severely at- 
tacked. It may also 
live saprophytically, and 
clusters of fruit-bodies 
are often found on old 
stumps and stems, on 
old timber, and in the 
rich soil of woods. The 
rhizomorpha, living un- 
derground, can extend 
for considerable dis- 
tances and infect the 
roots of neighbouring 
trees, and spreads in 
this way the diseases 
known as " Harzsticken " 
and "Bark -Canker," 
which are very destruc- 
tive to young trees. 


Fie. 174. Psalliota campestvls. A Tangential section of 
pileus showing lamellfe (!) B Portion of gill more highly 
magnified ; t trama ; hy hymenium with basidia and basidio- 
spores ; sh, sub-hymenial layer. C A portion of the same 
more highly magnified ; s' s" s'" s"" various stages in the 
development of basidiospores ; q paraphyses. 

The chief charac- 
teristics by which 
the numerous genera 
are separated are 
the presence or the absence of the two kinds of veils, the nature 
of the fruit-body, the form, branching of the lamella?, and their 
position and relation with respect to the stem, the shape of the 



pileus, the colour of the spores, etc., etc. A knowledge of the 
colour may be obtained by plac- 
ing the pileus with the lamellee 
turned downwards on a piece of 
white or coloured paper, so that 
the spores, as they fall off, are 
collected on the paper, and. the 
arrangement of the lamellae can 
then be clearly seen. 

FIG. 175. Canthardlus cibarius (reduced). 

FIG. 176. Armillaria mellca (| nat. sizel : 
a root of a Fir; b rhizomorpha-strands ; 
c-f fruit-bodies in four different stages of 

FIG. 177. The mycelium of Armillaria mellea (" Rhizomorplta") (nat. size). 



About 4,600 species belonging to this order have been described. 

On account of the large number of species the order is divided into several 
sections : 

1. Agaricinei ; fruit-body fleshy ; lamellas membranous, knife-like, with 
sharp edge ; basidia crowded together. The FOLLOWING HAVE WHITE SPORES : 
Amanita (Fly Mushroom), with volva, and generally also the upper ring on the 
stalk; many are poisonous, such as A. muscaria (Fig. 178) which has bright 
red pileus with white spots, A. pantherina and A.phalloides; A. ccesarea is edible. 
Lepiota procera (Parasol Fungus) is one of the largest Mushrooms ; it has a 
scaly pileus and moveable ring (edible). Armillaria mellea has been mentioned 
above (Figs. 176, 177). Tricholoma, lamellae indented near the stalk; T. gam- 
bosun (Pomona Fungus) belongs to the best of edible Fungi ; T. personatum 
often forms fairy rings 
(see above). Clitocybe, 
lamella decurrent ; C. 
nebularis is edible. 
Pleurotus, stalk eccen- 
tric ; P. ostreatus (Oyster 
Mushroom) grows in 
clusters on tree-stems 
(edible). Collybia and 
Myccna, species numer- 
ous, small. SPOKES 
ROSE-BED : Volvaria and 
Hypor hod ius. SPORES 
BROWN : Cortinariiis, 
with cob web -like veil ; 
Pholiota, membranous 
veil and ring ; P. squar- 
rosa in clusters on tree- 
stems ; P. mutabilis, on 
tree-stumps (edible). 
Hypholoma,Psalllota; to 
this section the common 
edible Mushroom (Fig. 

FIG. 178. Fly Mushroom (Amanita juwscaria). 

172-174) belongs, with annulus and chocolate-coloured lamellaa ; it is cultivated 
for the sake of the fine flavour. SPORES BLACK: Coprinarius. 

2. Marasmiei. Fruit-body tough, almost leathery, and persistent ; spores 
white. Marasmius oreades forms large, regular fairy-rings on pastures and 
commons ; it is used as seasoning in food. Panus stipticus with eccentrically- 
placed stalk, in clusters on tree-stumps. Schizophyllum has the edge of the 
lamellae divided longitudinally, and the split portions revolute. Lentinus 
affords a transition to Dcedalea among the Polyporacese. 

3. Russulei. Fruit-body fleshy and fragile, in which two different systems 
of hyphse may be distinguished ; spores thorny, white, or pale-yellow. Many are 
poisonous. Russula has generally fragile and thick lamellffi reaching from 
stalk to edge of pileus ; pileus frequently red. Lactarius has white or yellow 


milky juice, which often is very acid. L. deliciosus has red-yellow milky juice, 
and is of a pleasant flavour. L. torminosus is poisonous. 

4. Hygrophorei. Lamella thick and waxy, widely separated ; spores 
white. Many species of Hygrophorus have brightly-coloured pileus and grow 
among the grass on moors and commons. Nyctalis is parasitic on larger Toad- 
stools. It is remarkable for its abundant formation of chlamydospores, whilst 
the basidiospores are little developed. 

5. Coprinei. Fruit-bodies very soft, quickly perishable ; lamellae membran- 
ous and deliquescent. The basidia are separated from each other by para- 
physes. Coprinus has coal-black spores, grows on manure, and sometimes 
developes sclerotia. 

6. Paxillei. Fruit-body fleshy ; lamellrc easily detached from the pileus and 
reticulately-joined near the stalk. They form a connecting link between the 
Agaricacere and Boletus. 

7. Cantharellei. Lamellfe reduced to dichotomously-divided folds, decur- 
rent on the stalk. Cantharellus cibarius (Fig. 175) is yolk-yellow, and grows on 
the ground in woods (edible). It is allied to Craterellus. 

Family 3. Phalloidese. 

The fruit-bodies before they are ripe are spherical or ovoid, and 
enclosed by a fleslty covering, the peridium, which, is perforated at 
maturity and remains as a sheath (Fig. 179) ; the fruit-bodies 
are hemiangiocarpic. 

Order 1. Phallacese (Stink-horns). The peridium has a 
complicated structure and is composed of three layers, the in- 
termediate one being thick and gelatinous. The gleba (the tissue 
which bears the hymenium) is situated upon a peculiar receptacle 
which expands into a porous stalk and by its sudden distension, 
rupturing the peridium, elevates the gleba and hymenium above 
the peridium, which remains as a sheath. The gleba becomes 
gelatinous and dissolves away as drops. To this order belong many 
peculiar and often brightly coloured forms, which are natives of 
the Southern Hemisphere. 

Phallus impudicus (Stink-horn) (Fig. 179), has a fruit-body which at first is 
white, heavy, and soft, and resembles a hen's egg in shape and size. The peri- 
dium is divided into three layers (Fig. 179 e, g,f) of which the external and 
internal are membranous, and the middle one very thick and gelatinous ; each 
of these has again a laminated structure. The peridium when ruptured re- 
mains as a sheath (k) at the base of the stalk. The receptacle at first is 
strongly compressed (h) but afterwards expands into a long stalk (I) which 
bears the conical gleba (?). Prior to the rupture of the peridium the gleba 
consists of a greenish mass ((') which, when exposed, emits a carrion-like stench 
serving to attract flies, by whose agency the spores are distributed. It is found 
commonly in hedgerows and in woods, growing on the ground. The much 



smaller and less common P. caninus is found on rotten tree-stumps. In 
Clathrus cancellatus the receptacle expands into a bright red, reticulate struc- 
ture. A native of the South of Europe. Coins, Aseroe, Mitromyces. 

Order 2. Sphaerobolacese. An intermediate layer of the peridium swells 
when ripe, becomes convex, and ejects the remaining spherical portion of the 
fruit-body which contains the spores. Sphcerobolus carpobolus has small, 
spherical fruit-bodies which open in the form of a star. 

FIG. 179. Phallus tmpiidicus (Stink-horn), somewhat diminished. Fruit-bodies in all 
stages of development (b, c, d and 7c-m) are seen arising from a root-like mycelium (a) ; d 
longitudinal sections through a fruit-body b3fore the covering has ruptured. 

*Family 4. Gasteromycetes. 

The fruit-body is angiocarpic, fleshy at first, and later generally 
more or less hard and continues closed after the spores are ripe. 
The tissue lying immediately inside the peridium is termed the 
yleba ; it is porous, containing a larger or smaller number of 
chambers lined with the hymenium, which is either a continuous 



layer of basidia or else it fills up the entire cavity. The basidia 
as a rule bear four spores, sometimes eight (Geaster), or two 
(Si/menogaster). The tissue of the walls (trama) consists often 
(Lycoperdacece} of two kinds of hyphre, some thin and rich in 
protoplasm, divided by transverse septa and bearing the basidia ; 
others thicker and thick-walled which do not dissolve like the 
former on the ripening of the spores, but continue to grow and 
form a woolly, elastic mass, the capillitium, which may be regarded 
as highly developed paraphyses. The peridium may be either 
single or double, and presents many variations in its structure 
and dehiscence. The mycelium is generally a number of string- 
like strands, living in soils rich in humus. 

Order 1. Tylostomaceae. Capillitium present. After the rupture of the 
peridium the remaining part of the fruit-body is elevated oil a long stalk. 
Tylostoma inainmosum, on heaths. 

Order 2. Lycoperdaceae. The fruit-body has a double 
peridium ; the external one at length breaks into fragments 
(Lycoperdon, Bovista), or it has a compound structure of several 
layers (Geaster) and detaches itself as a continuous envelope from 
the inner layer, which is membranous and opens at its apex. 
The interior of the fruit-body consists either solely of the fertile 
gleba (Bovista, Geaster), or, in addition, of a sterile tissue at the 
base (Lycoperdon). A capillitium is also present. 

Lycoperdon (Puff-ball) has a sterile part at the base of the fruit-body which 
often forms a thick stalk. The surface of the peridium is generally covered 

with warts or projections. When 
young this Fungus is edible, but 
when ripe it is dry, and used for stop- 
ping the flow of blood. L. giganteum, 
which is often found growing in 
meadows, attains a considerable size, 
its diameter reaching as much as 
eighteen inches. L. gemmatum (Fig. 
180) is covered with pyramidal warts ; 
in woods. Bovista has no sterile 
basal part; the external peridium is 
smooth, and falls away in irregular 
patches. B. plumbea, on linka near 
the sea. Geaster (Earth- star) has an 
external peridium composed of several 
layers, which when the fruit-body opens, split into several stellate segments. 
These segments are very hygroscopic, and in dry weather bend backwards and so 
raise the inner peridium into the air. The inner peridium contains the spores. 

FIG. 180.- 

' rdon gemmutum 
(a nat. size). 



and capillitia. G. coliformis has several apertures in the inner peridium. The 
other species have only one regular aperture at the apex. G. striatits has a 
pedicellate inner peridium, with conical, striped peristome. G. fornicatus has 
an external peridium split into four segments. This last and several other 
species produce " mycorhiza " on the roots of Conifers. 


FIG. 181. I Hynienogaster c'drimis (nat. size) ; II longitudinal section through H. tener 
( x 5) ; III portion of a section of H. caloxporus ; g a chamber ; h hymenium ; sp. spores; 
t trama (x 178) ; IV Rliizopogon litieulus (nat. size); V Scleroderma vulgare, VI section of V ; 
VII basidia with spores belonging to the same Fungus. 

Order 3. Sclerodermatacese. Capillitium wanting. The 
peridium is simple and thick, gleba with round, closed cham- 
bers, which are filled with basidia. 

Scleroderma has a corky peridium. The fruit-bodies commence their develop- 
ment under ground. S. vulgare (Fig. 181 V-YIIJ, has a hard, slaty-black gleba. 


Order 4. Nidulariaceae (Nest-Fungi). Small Fungi of 
which the fruit-body at first is spherical or cylindrical but upon 
maturity it becomes cupular or vase-like, and contains several 
lenticular " peridiola " lying like eggs in a, nest. The peridiola 
are the chambers which contain the hymenium, covered by a thin 
layer of the gleba, all the remaining portion of the gleba becoming 
dissolved. On decaying wood. 

Nidularia has spherical fruit-bodies containing a large number of lenticular 
peridiola, embedded in a slimy mass. Crncibuhim has fruit-bodies resembling 
crucibles with discoid peridiola, each with a spirally-twisted stalk. Cyathus 
has a fruit-body, which when open is campauulate, with stratified peridium, 
and long-stalked, lense-shaped peridiola. 

Order 5. Hymenogastraceae. Fruit-bodies tubercular, 
globose and subterranean, resembling very closely the Truffles, 
from which they can only be distinguished with certainty by 
microscopic means. The peridium is simple, capillitium wanting, 
and the gleba encloses a system of labyrinthine passages covered 
with a continuous hymenium. The fruit-bodies persist for some 
time, and form a fleshy mass, the spores being only set free by 
the decay of the fruit-body, or when it is eaten by animals. The 
majority are South European. Hymenogaster, ^lelanoijaster, 
Rhizopoijon (Fig. 181 I-IV). 

Basidiolichenes (Lichen-forming Basidiomycetes). 

Several Fungi belonging to the Basidiomycetes have a symbiotic 
relationship with Alga? exactly similar to that enjoyed by certain 
Ascomycetes, and these are therefore included under the term. 
Lichens (p. 136). They are chiefly tropical. 

Order 1. Hymenolichenes. To this order belong some 
gymnoca.rpic forms : Cora, Dictyonema, Laudatea. l 

Order 2. Gasterolichenes. To this belong some angiocarpic 
forms : Emericella, Trichocoma. 

Fungi imperfect! (Incompletely known Fungi). 

1. The Saccharomyces-forms are Fungi which are only 
known in their yeast-conidial form. They are conidia of higher 

1 The two last genera are identical, the Algal part being a Scytonema, that of 
Cora a Chroococcus ; while the same Fungus a Thelephora takes part in the 
formation of all three (A. Holler, Flora, 1893). 



Fungi which can multiply to an unlimited extent by budding 1 in 
nutritive solutions, and in this way maintain their definite size and 
shape. The budding takes place only at the ends of the conidia. 
The wall of the conidium forms at one or at both ends a small wart- 
like outgrowth, which gradually becomes larger, and is finally 
separated from its mother-cell as an independent cell, surrounded 
by a closed cell- wall (Fig. 182 a, b). 

FIG. 182. Beer-yeast (Saccharomyces cerevisice) : a-b (x 400); c-f ( x 750) ; c a cell in the 
process of forming spores ; d a cell with four ripe spores; e the spores liberated by the 
dissolution of the cell-wall ; / three germinating spores ; g mycelium-like cell-chains. 
(x 1030: after Em. Chr. Hansen.) 

Under very favourable conditions multiplication occurs so 
rapidly that the daughter-cells themselves commence to form 
buds, before they have separated from their mother-cell, with the 
result that pearl-like chains of cells are produced. When the 
yeast-cells have only limited nutriment, with an. abundant supply 

w. B. N 


of air, at a suitable temperature, an endogenous formation of 
spores takes place. The protoplasm of the cells divides into 1-4 
(rarely a greater number) masses (Fig. 182 c, d, e) which surround 
themselves with a thick cell- wall, and in this state can with- 
stand adverse conditions and periods of dryness lasting for several 

The sporangia are not asci since they have no definite form, and 
a definite number, form and size of spores is not found. The 
spores in the different species and kinds occupy varying periods 
for their development, although exposed to the same temperature, 
a fact of importance in determining one from another. On 
germination the wall of the mother-cell is destroyed, and each 
spore gives rise to a new cell, multiplication taking place by 

budding (Fig. 182 /). The majority 
of Yeast-Fungi are able to produce 
alcoholic fermentation in saccharine 

The most important of these 
Fungi is the Beer-yeast (Saccharo- 
mijcescerevisice) with ovate, ellipsoidal 
or spherical cells (Fig. 182). It is 
a plant which has been cultivated 
from time immemorial, on account 
of its property of producing alco- 
holic fermentation in sugar-contain- 
ing extracts (wort), derived from 

germinating barley (raalt ). C ar- 

bonic acid is also set free during this process. The " surface- 
yeast " (Fig. 182 a), which produces ordinary beer when the 
brewing takes place at higher temperatures, has cell-chains ; 
" sedimentary yeast " (Fig. 182 b), used in the brewing of Bavarian 
beer, has spherical cells, solitary, or united in pairs. Both these 
and the following Yeast-Fungi include, according to Hansen, 
several species and kinds. 

The " Ferment of Wine " (Saccharomyces ellipsoideus) produces 
wine in the juice of grapes. Uncultivated yeast-cells are always 
present on grapes ; an addition of this species to the " must " is 
hot necessary to secure fermentation. A large number of other 
" uncultivated " yeast-cells appear in breweries mixed with 
the cultivated ones, and cause different tastes to the beer (S. 
pastorianus, etc.). S. ludwigii, found, for instance, on the slimy 


discharge from Oaks, produces abundant cell-chains on cultiva- 
tion. S. apiculatus is very frequently met with on all kinds of 
-sweet fruits, it has orange-like cells. S. mycoderma has cylindri- 
cal cells, often united together in chains (Fig. 183) : it forms a 
-whitish-gray mass (" fleur de vin ") on wine, beer, fruit-juice, etc., 
standing- in bottles uncorked or not entirely filled. It is thought 
that this Fungus causes decomposition and oxydises the fluid in 
which it is found, but it cannot produce alcoholic fermentation in 
saccharine liquids, and it does not form endospores ; hence it is 
uncertain whether it is true Saccharomyces. 

The "Dry-yeast" used in baking white bread is "surface- 

FIG. 184. Oidiitm lactts : a branched hypha commonly met with; b a hypha lying in 
milk and producing aerial hyphae which give rise to oidia ; c a branch giving 1 rise to 
oidia, the oldest (outermost) oidia are becoming detached from one another ; d a chain 
of divided cells; germinating oidia in different stages (slightly more magnified than the 
other figures). 

yeast." In leaven, a kneaded mixture of meal, barm and water, 
which is used for the manufacture of black bread, Saccharomyces 
minor is present, and a species allied to this produces alcoholic 
fermentation in dough with the evolution of carbonic acid, which 
causes the dough to " rise." 

2. Oidium-forms. Of many Fungi only the Oidium-forms 
are known, which multiply in endless series without employing 
any higher form of reproduction. Oidittm lactis (Fig. 184) is an 
imperfectly developed form which frequently appears on sour 


milk and cheese. It can produce a feeble alcoholic fermentation 
in saccharine liquids. Thrush or aphthaB (0. albicans) appears as 
white spots in the mouths of children. Several similar Oidium- 
forms are parasites on the skin and hair of human beings, and 
produce skin diseases, such as scurvy (0. sclwenleinii) and ring- 
worm (0. tonsurans)i 

3. Mycorhiza. These Fungi, which have been found on the 
roots of many trees and heath-plants, particularly Cupuliferse and 
Ericacea?, consist of septate hyphse, and belong partly to the 
Hymenomycetes, partly to the Gasteromycetes. It has been 
shown that the Mycorhiza enters into a symbiotic relationship 
with the roots of higher plants. 



In this Division a well-marked alternation of generations is to be 
found. The development of the first or sexual generation (gameto- 
pliyte), 1 which bears the sexual organs, antheridia and archegonia, 
commences with the germination of the spore, and consists, in the 
Liverworts, of a thallus, but in the true Mosses of a filamentous 
protonema, from which the Moss-plant arises as a lateral bud. 
The second or asexual generation (sporophyte), developed from the 
fertilised oosphere, consists of a sporangium and stalk. 

The sexual generation, the gametophyte. The protonema 
in the Liverwoi-ts is very insignificant, and not always very sharply 
demarcated from the more highly developed parts of the nutritive 
system. In the true Mosses the protoiiema is well-developed, and 
consists of a branched, alga-like filament of cells, the dividing 
cell-walls being always placed obliquely. In the parts exposed to 
the light it is green, but colourless or brownish in those parts 
which are underground (Fig. 186). The protonema is considered 
to be a lower form of the stem, and grows in the same manner 
by means of an apical cell ; at its apex it may directly develope 
into a leaf-bearing stem, or these arise from it as lateral branches 
(Fig. 186 it). 

The more highly differentiated part of the vegetative system, 
the " Moss-plant," which is thus developed from the protonema, 
is in the " thalloid " Liverworts generally a dichotomously- 
branched thallus without any trace of leaf-structures (Fig. 
194) ; in Marchantia (Fig. 197) and others, scale-like leaves 
(amphigastria) are found on the under surface. The higher 
Liverworts and the Leafy-Mosses are differentiated into a fila- 
mentous, ramified stem with distinct leaves arranged in a definite 
manner, resembling the stem and leaves of the higher plants (Figs. 
186, 195, 200). 

1 Formerly termed oophyte. 



True roots are wanting, but are biologically replaced by rhizoids. 
These are developed on the stems or thallus : in the Liverworts 
they are unicellular, but in the Leafy-Mosses generally nralticel- 
lular and branched. In the latter group they are considered 
identical with the protonema, and may become true protonema, 
and new plants may be developed from them (Fig. 186 6). 

The internal structure of the sexual generation is very simple. 
The leaves in nearly all cases are formed of a single-layered plate 
of cells ; in the Leafy -Mosses, however, a midrib is very often 

FIG. 186. A Lower portion of a Moss-plant with rhizoids (r), one of which bears a 
reproductive bud (l>). The dotted line indicates the surface of the ground ; the portions 
projecting above this become green protonema (p); fc is a young Moss-plant formed on 
one of these. B Germatining spore of Funaria hygrometrica, with exospore still 
attached. C, D Older stages of the protonema. 

formed, and sometimes, also, marginal veins ; and along these lines 
the leaves are several layers of cells in thickness. The stem is 
constructed of cells longitudinally elongated, the external ones of 
which are narrower and sometimes have thicker walls than the. 
more central ones. Vessels are not found, but in several Mosses 
there is in the centre of the stem a conducting strand of narrow, 
longitudinal cells, which represents the vascular bundle in its first 


stage of development. This strand contains elements for convey- 
ing water as well as sieve-tubes. Stomata are entirely wanting in. 
the sexual generation of the Leafy-Mosses ; they are found in a 
few Liverworts (Marchantia) ^ but their structure is not the same 
as in the higher plants. 

VEGETATIVE REPRODUCTION takes place by gemmae or buds which 
arise on the protenema, the rhizoids, the thallus, or the shoots, and 
become detached from the mother-plant ; or else the protonema 
and the older parts of the plant simply die off, and their branches 
thus become independent plants. This well-developed vegetative 
reproduction explains why so many Mosses grow gregariously. 
In certain Marchantiacea? special cupules, in which gemmre are 
developed, are found on the surface of the thallus (Fig. 197 J,s-,*). 

FIG. 137. Marchantia polymorpha : FIG. 188. Spermatozoids. 

a mature untheridium. 

Again, protonema may also arise from the leaves, and thus the 
leaves may act as reproductive bodies. Certain Mosses nearly 
always reproduce vegetatively, and in these species the oospheres 
are seldom fertilised. 

The first generation bears the SEXUAL ORGANS ; both kinds are 
found either on the same plant (monoecious), or on separate plants 
(dioecious). In the thalloid Liverworts they are often situated on 
the apex of small stems (gametophores'), springing from the surface 
of the thallus. In the Leafy- Liverworts and true Mosses the leaves 
which enclose the sexual organs often assume a peculiar shape, and 
are arranged more closely than the other leaves to form the so-called 
" Moss-flower." The male sexual organs are called antheridia. 
They are stalked, spheroid, club- or egg-shaped bodies whose 
walls are formed of one layer of cells (Fig. 187), enclosing a mass 
of minute cubical cells, each one of which is a mother-cell of 


ti spermatozoid. The spermatozoids are self-motile ; they are 
slightly twisted, with two cilia placed anteriorly (Fig. 188), while 
posteriorly they are generally a trifle club-shaped, and often bear 
at that part the remains of the cytoplasm, the spenuatozoid itself 
being formed from the nucleus. In the presence of water the ripe 
antheridinm bursts, and its contents are ejected ; the spermato- 
zoids, being liberated from their mother-cells, swarm about in the 
water in order to effect fertilisation. 

The female sexual organs are termed archegonia. They are flask- 
shaped bodies (Fig. 189), the lower, swollen portion (venter) having 

FIG. 189. Marchantia polyrnorpha. A A young, and B a ripe archegonium with open 
neck. C An unripe sporangium enclosed by the arcaegonium a : st the stalk ; / the wall 
of the sporangium. Elaters are seen between the rows of spores. 

a wall, in most cases from 1-2 cells thick, enclosing the oosphere 
(Fig. 189 B, Z.-) : the long neck is formed of tiers of 4-6 cells, 
enclosing a central row of cells the neck-canal-cells (Fig. 189 A). 
When the archegonium is fully developed, the walls of the neck- 
canal-cells become mucilaginous and force open the neck of the 
archegonium. The mucilage thus escapes, and, remaining at the 
mouth of the archegonium, acts in a somewhat similar manner to 
the stigma and conducting tissue of a carpel, by catching and con- 
ducting the spermatozoids to the oosphere (Fig. 189 B, m), with 



whose cell-nucleus they coalesce. With regard to the formation 
of the oosphere, it may further be remarked that the lower part of 
the archegonium originally encloses the so-called " central cell "; 
but shortly before the archegonium is ripe, this cuts off a small 

portion, the ventral- canal- cell, which 
lies immediately beneath the neck, 
and the larger, lower portion be- 
comes the oosphere. 

The organs mentioned here, antheridia 
and archegonia, are present in the Crypto- 
gams (Pteridophyta) and the Gymno- 
sperms. They have always the same 
fundamental structure, but with slight 
modifications of detail. These plants are 
therefore known as the AIICHEGONIATA.. 

The fertilisation of the Mosses 
cannot be effected without "water. 
Rain and dew therefore play a 
very important part in this pro- 

FIG. 190. Andrecea rupestris. Longi- 
tudinal section through a sporangium 
at the time when the mother-cells of 
the spores are dividing : p pseudo- 
podium ; / foot ; D vaginula ; h neck ; 
c columella ; w wall of the sporan- 
gium ; e external row of cells; s the 
spore-sac ; t the spore-mother-cells ; 
r the calyptra with the neck of arche- 
gonium (zj. 

FIG. 191. Andreaa rupestris. Transverse 
section through a ripe sporangium. In the 
middle is seen the four-sided columella, sur- 
rounded by the numerous spores, drawn dia- 
grammatically. Surrounding them is seen the 
wall of the sporangium, whose outer layer of 
cells is thickened and coloured. The layer of 
cells is unthickened in four places (x), indi- 
cating the position of the clefts (see Fig. 

cess, and for this end various modifications of structure are found. 


Among the sexual organs, paraphyses filamentous or club- 
shaped bodies are to be found. 

The asexual generation, the sporophyte (Moss-fruit or 
sporogonium); As the result of fertilisation the oosphere sur- 
rounds itself with a cell- wall, and then commences to divide in 
accordance with definite laws. 1 The embryo (Fig. 189 G) pro- 
duced by these divisions remains inside the wall a-a of the arche- 
gonium (Figs. 190, 199 D, E), and developes into the sporogonium, 
which remains attached to the mother- plant, often nourished by 
it, as if the two were one organism. The lower extremity of the 
sporogonium, the foot (Figs. 190 /; 199 _D), very often forces its 
way deep down into the tissue of the mother-plant, but without an 
actual union taking place. The central portion of the sporogonium 
becomes a shorter or longer stalk (seta), while the sporangium 
itself is developed at the summit. At a later stage, during the 
formation of the spores, the sporangium very often assumes the 
form of a capsule, and dehisces in several ways characteristic of the 
various genera (Figs. 192, 193, 194, 195, 200). The basal portion 
of the archegoninrn grows for a longer or shorter period, forming 
a sheath, the calyptra, in which the capsule is developed, but 
eventually it ceases to enlarge, and is then ruptured in different 
ways, but, quite characteristically, in each group. Anatomically, 
the asexual generation is often more highly differentiated than the 
sexual ; thus, for instance, stomata are present on the sporangia 
of the true Mosses, but are absent in the sexual generation. 

As the capsule developes, an external layer of cells the amphi- 
thecium and an internal mass the endothecium are differ- 
entiated. As a rule the former becomes the wall of the capsule 
while the latter gives rise to the spores. In this Division, as in 
the Pteridophyta, the name archesporium (Fig. 190 t) is given to 
the group of cells inside the sporangium which gives rise to the 
mother-cells of the spores. The archesporium is in general a 
unicellular layer ; in Spliaijnum and Anthoceros it is derived from 
the most internal layer of the amphithecium, but with these ex- 
ceptions it arises from the endothecium, usually from its most 
external layer. In the true Mosses and in liiccia only spore- 
mother-cells are produced from the archesporium, but in the 

1 The oospore divides by a wall transverse or oblique to the longer axis of 
the archegoniuni. From the upper (epibasal) cell, the capsule (and seta) is 
derived, while the lower (hypobasal) gives rise to the/oo(. In Riccia the hypo- 
basal half takes part in the formation of the sporangium. 


majority of the Liverworts some of these cells are sterile and 
become elaters (cells with spirally thickened walls, Figs. 196, 189), 
or serve as "nurse-cells" for the spore-mother-cells, which gradu- 
ally absorb the nutriment which has been accumulated in them. 
In Anthoceros, and almost all the Leafy-Mosses, a certain mass of 
cells in the centre of the sporangium (derived from the endo- 
thecium) does not take 
part in the formation 
of the archesporium, 
but forms the so called 
"column" or " colum- 
ella" (Figs. 190,191). 
The spores arise in 
tetrads, i.e. four in each 
mother-cell, and are 
arranged at the corners 
of a tetrahedron, each 
tetrahedron assuming 
the form of a sphere or 
a triangular pyramid. 
The mature spore is a 
nucleated mass of pro- 
toplasm, with starch or 
oil as reserve material. 
The wall is divided into 
two layers: the exter- 
nal coat (exospore) 
which is cuticularized 
and in most cases 
coloured (brown, yel- 
lowish), and the internal 
coat (endospore), which 
is colourless and not 
cuticularized. On ger- 
mination the exospore 
is thrown off, the endo- 
spore protrudes, and 
cell-division commences 
and continues with the growth of the protonema (Fig. 186, B-D). 

The morphological explanation which Celakovsky has given of the sporo- 
gonium, and which is not at all improbable, is, that it is homologous with 

FIG. 193. Andrecea petro- 
phila. An empty capsule ; 
the calyptra has fallen off. 
(Mag. 25 times.) 

FIG. 192. Andreaa petropJiila. A ripe sporogoninm : 
a an archegonium which has been raised with the 
pseudopodium ; p the foot ; 6 the neck ; d-e the dark- 
coloured portion of the sporangium, whose outer cell- 
walls are considerably thickened ; c-c the thin-walled 
portions where the dehiscence occurs ; o the lower ex- 
tremity of the spore-sac; /calyptra; g the apex of 
the sponmgium. (Mag. 25 times.) 


an embryo consisting of a very small stem-portion and a terminal spore-pro- 
ducing leaf. This will be further explained in the introduction to the Flower- 
ing-plants (p. 236). 

In the Liverworts the young sporogouium lives like a parasite, 
being nourished by the sexual generation (only in Anthoceros has it 
a slight power of assimilation). In the Leafy- Mosses, on the other 
hand, with regard to the power of assimilation, all transitions 
are found from abundant assimilation (Funaria, Physcomitrium') 
to almost complete " parasitism " (Sphagnum, Andrecea). In the 
majority of the operculate Mosses the sporogonium has a more 
or less perfect system of assimilation, and is able itself to form a 
large portion of the material necessary for the development of the 
spores, so that it chiefly receives from the sexual generation the 
inorganic substances which must be obtained from the soil. The 
more highly developed the assimilative system of the sporogonium, 
the more stomata are present. 

APOSPORT. In some operculate Mosses it has been possible to obtaiu a pro- 
tonema with small Moss-plants from the seta, when severed from its Moss- 
pls,nt, and grown on damp sand. 

The Mosses are the lowest plants which are provided with stem 
and leaf. They are assigned a lower place when compared with 
the higher Cryptogams, partly because there are still found within 
the Division so many forms with a mere thallus, partly because 
typical roots are wanting and the anatomical structure is so ex- 
tremely simple, and partly also because of the relation between 
the two generations. The highest Mosses terminate the Division, 
the Muscineas and Pteridophyta having had a common origin in 
the Algae-like Thallophyta. 

They are divided into two classes : 

HEPATIC^;, or Liverworts. 

Musci FKONDOSI. True Mosses or Leafy-Mosses. 

Class i. Hepaticae (Liverworts). 

The protonema is only slightly developed. The remaining part 
of the vegetative body is either a prostrate, often dichotomously- 
braiiched thallus, pressed to the substratum (thalloid Liverworts), 
with or without scales on the under side (Figs. 194, 197) ; or a 
thin, prostrate, creeping stem, with distinctly-developed leaves, 
which are borne in two or three rows (Figs. 195, 198), viz., two on 
the upper and, in most cases, one on the under side. The leaves 
situated on the ventral side (amphigastria) are differently shaped 
from the others (Fig. 198 a), and are sometimes entirely absent. 



In contradistinction to the Leafy-Mosses, stress must be laid on 
the well-marked dorsivenlrality of the vegetative organs ; i.e. the 
very distinct contrast between the dorsal side exposed to the light 
and the ventral side turned to the ground. Veins are never found 
in the leaves. 

The ventral part of the archegonium (calyptra) continues to grow 
for some time, and encloses the growing embryo, but when the 
spores are ripe it is finally ruptured by the sporangium, and re- 
mains situated like a sheath (vaginula) around its base. The 
sporangium opens, longitudinally, by valves or teeth (Fig. 194, 195, 
197 6), very rarely by a lid, or sometimes not at all. A columella 
is wanting (except in Anthoceros, Fig. 194) ; but on the other hand, 
a few of the cells lying between the spores are developed into 

PIG. 194. Anihoceros Iwis 
(nat.size): K-K capsules. 

FIG. 196. An 
elater with two 


FIG. 195. PlagiocJiila asplenioides : a unripe, and b an open capsule; p involucre. The 
ventral edge of each leaf is higher than its dorsal edge, and covered by the dorsal edge of 
the next one. 

slaters (Fig. 196), i.e. spindle-shaped cells with spirally-twisted 
thickenings, which are hygroscopic, and thus serve to distribute 
the spores. (They are seen in Fig. 189 C, not yet fully developed, 
as long cells radiating from the base of the sporangium. They 
are wanting in Riccia). 

Round the entire archegoniurn, (or group of archegonia, when several are 
developed on the same receptacle) a sheath the involucre is often formed, 
which persists, and encloses the base of the stalk of the sporangium, together 
with the sheath of the archegouium (Fig. 195 p). In the Marchantiaceae each 
archegonium is enclosed in a loose investment, the perigyuiuin, which is 
developed as an outgrowth from the cells of its stalk. 



The majority of the Liverworts are found in damp and shady 
places, pressed to the substratum ; a few are found floating in 
fresh water. 

Family 1. Marchantieae. 

This embraces only forms with a thallus, which is more or less 
distinctly dichotomously branched, in some, one or two rows of 
thin leaves are situated on its under surface. On the upper 
surface of the thallus are found large air-chambers. 

Order 1. Ricciacese. The sporogonia are, with the excep- 
tion of a few genera, situated singly on the surface of the thallus, 
and consist only of a capsule without foot or stalk. They always 
remain enclosed by the wall of the archegonium (calyptra), and open 


FIG. 197. Narchantla, polymorpha. A Female plant (nat. size) : a and b are arche- 
goniophores in various stages of development ; s cupules with gemmse (see page 183). 
B An archegoniophore seen from below, the short-stalked sporangia are seen placed in 
8-10 double rows. C Male plant, with a young and an older antheridiophore. D 
Antheridiophore halved vertically to show the antheridia (h) ; in the aperture of the 
pits in which they are sunk the older ones to the left, the younger to the right. 

only by its dissolution. Elaters are not developed. Some genera 
are found floating like Duckweed. Riccia glauca grows on damp 
clay soil. _K. JJaitans and R. natans float in stagnant waters. 

Order 2. Corsiniaceae. (Not native). Intermediate forms between the 
preceding and the following order. In internal and external structure mainly 
resembling the Marchantiaceffi. Comma ; Boschia. 

Order 3. Marchantiacese, are large, fleshy forms. The 



surface of the thallus is divided into small rhombic areas, in the 
centre of each of which is found a large, peculiarly constructed 
stoma (Fig. 197 A) ; beneath each of these a large air-cavity is to 
be found. From the floor of the air-cavity a number of alga- 
like cells project into it ; these contain chlorophyll and are 
therefore the assimilating cells. The antheridia and archegonia 
are each found aggregated on specially formed branches (some- 
what resembling Mushrooms) projecting from the surface of the 
thallus. The antheridia are developed on the upper surface (Fig. 
197 (7, D) and the archegonia on the lower (Fig. 197 A, B), near 
the centrally-placed stalk. 

Mar chant ia poly morpha is dioecious (Fig. 197), and very common 
on damp places. Lunularia (South Europe), frequently found 
on flower-pots in conservatories ; Preissia, Fegatella, Reboulia, 

Family 2. Anthoceroteae. 

These have an entirely leafless, fleshy, flat, and irregularly-shaped thallus. 
In its intercellular chambers Nostoc-colonies are often found, which have forced 
their way through the stomata situated on the under side. The antheridia an'd 
archegonia arise from the cells lying inside the thallus. The capsule resembles 
a long, thin pod; it has two valves and a columella. Anthoceros (A. he a is, 
Eig. 194, and punctatus). 

Family 3. Jungermanniese. 

Some forms in this family have a thallus in which leaf-like 
structures are found (Blasia), while in 
others (e.g. Metzgeria, Pellia, Aneura) they 
are entirely absent. The majority, how- 
ever, have round, thick stems, bearing 
dorsally two rows of leaves, and one row 
ventrally. Some of these have the leaves 
"underlying" (Fig. 195), while in others 
(Fig. 198) they are " overlying." (See Figs. 
195, 198, with explanation). 

The sporangia are spherical, stalked, and 
situated singly on the apex of the branches, 
and open by four valves (in Sphcerocarpus 
they are indehiscent). 

All the species in this family were 
formerly reckoned as belonging to one 
genus, Jungermannia, but now they are 
divided into several, arranged as follows : 

FIG. 198. Frullania dila- 
tata. Portion of a branch 
seen from the under side : 
v and b are the anterior and 
posterior edges of the same 
dorsal leaf; a ventral leaves 
(amphigastria). The dorsal 
leaves are " overlying," i'.e. 
the anterior edge of the 
leaf overlaps tLe posterior 
edge of the preceding one. 

192 f MUSCINEJ;. 

I. ANACROGYX.E. The archegonia are situated on the upper 
side of the thallus or stem, placed laterally, and covered by an 
" involucre," formed by the calyptra together with the tissue of 
the stem or thallus. 

a. ANELATERE.E. Without any elaters : Sphcerocarpus, Eiella. 

b. ELATEREJ;. a. Thalloid : Aneura pinguis, in damp situations ; 
Metzgeria furcata. on trees; Pellia epipliylla, in damp situations; 
Blasia pusilla, on damp clay soil, in the shade (scales are present 
on the thallus). ft. Foliose and ot dorsi ventral : Haplomitrium 

II. ACROGYNJ;. The apex of the stem or of certain branches is 
adapted for the formation of female shoots. The archegonia are 
most frequently aggregated on the apex of the shoots, and are 
encircled by their leaves (perichgetium). Between these and the 
archegonia, enclosing the latter, a peculiar cup-shaped organ (the 
involucre) is formed. This group only includes leaf-bearing 
genera : Frullania, Radula, Madotheca, Ptilidium, Calypogeia, 
Lepidozia, Mastigobryum, Lophocolea, Jungermannia, Scapania, 

Class 2. Musci frondosi or veri (True Mosses). 

In this class the protonema is well developed, and resembles a 
branched filamentous Alga, from which it can be easily distin- 
guished by its oblique septa (in Sphagnum it is a cellular expan- 
sion). The Moss-plant, which is developed directly from the 
protonema, generally has an erect, thick, cylindrical stem similarly 
constructed on all sides. The leaves are arranged spirally, the 
most frequent divergence being f or f (Fig. 200 A). A midrib is 
often present and also marginal veins formed by longitudinally 
elongated cells ; at these veins the leaf is more than one layer in 
thickness. In Leiicobryum the leaves are generally constructed 
of more than one layer. 

The stem grows by means of a three-sided, pyramidal, apical cell 
which gives rise to three rows of segments, each segment forming 
a leaf. The lateral branches arise from the lower portions of the 
segments, the upper portion of which does not take any part in the 
construction of the leaf. From their mode of origin the branches 
are not axillary, and differ in this respect from the Flowering- 

The ventral portion of the archegonium is very early ruptured 

MCSCINE5;. 193 

at its base by the growing sporogonium, upon which it remains, and 
it is thus raised into the air, forming a " hood," the caljptra 
(Figs. 192; 200 B). In the Sphagnacete the hood is not present; 
in this order, as in the Liverworts, the archegonium remains at the 
base of the sporogonium. The sporangium opens by circumsessile 
dehiscence ; the upper portion (operculum) being separated along 
a specially constructed ring of cells, and falls oft' like a " lid ' : 
(Fig. 200). Only in a few forms (families 2 and 3) does any vari- 
ation of this take place. Elaters are never found, but (with the 
exception of Archidium) there is always present in the sporangium 
a central mass of cells, the columella, which take no part in the 
formation of the spores. The columella, in some, does not reach 
quite to the operculum and in these cases the spore-sac is bell- 
shaped and covers the columella (Andrecea, Fig. 190 ; Sphagnum, 
Fig. 199 D) ; but in the majority of Mosses the columella extends 
to the lid, so that the space containing the spores becomes a 
hollow cylinder. 

The sporangium is generally raised on a long stalk ; in the great 
majority this stalk is formed from the lower half of the oospore 
and belongs to the asexual generation it is then known as the 
seta. In Andreaea and Sphagmim the seta is very short, and the 
sporangia are raised upon a long stalk (pseudopodium) developed 
from the summit of the sexual generation (Figs. 190, 192). In 
the latter figure an archegonium (a) is seen attached to the 
pseudopodium, having been carried up with this during the course 
of its development. The summit of the pseudopodium is enlarged 
to embrace the foot of the sporogonium (Figs. 192, 199 D) 

A. The sporangium is supported on a pseudopodium ; the columella does 
not extend to the operculum. 

Family 1. Sphagneae (Bog- Mosses). 

The protonema has been already described. The stem is 
regularly branched owing to the fact that a branch, or collection of 
branches, arises at every fourth leaf. These branches are closely 
covered with leaves, some are erect, while others hang down and 
surround the stem. No rhizoids are developed. These Mosses are 
of a whitish-green colour, and when water is present are always 
saturated with it like a sponge, the reason for this being found in 
the construction of the stem and leaves. The stems are covered 
by an external layer of large clear cells, without chlorophyll, but 
with annular or spiral thickenings on the walls, which are also 

w. B. 


perforated by large holes. By means of capillary attraction, water 
is thus raised to the summit of the stem. Similarly constructed 
cells are also found in the leaves, but they are sui'rounded by a net 
of very narrow, chlorophyll-containing cells (Fig. 199 F), whose 

colour is thus to a great extent lost 


those which are 

FIG. 199. Sphagnum acutifolium. A The upper portion of a plant : a branches with 
antheridia ; cJv branches with terminal archegonia and perichaetia; b the upper stem- 
leaves. B A male branch whose leaves are partly taken off in order to show the 
antheridia. C Group of three archegonia : the central one (a) is formed from the apical 
cell. D Sporogonium in longitudinal section : the broad foot (sg') is sunk in the vaginula, 
D ; c calyptra; or neck of the archegonium ; ps pseudopodium. E ripe sporangium with 
operculum, and the remains of the archegonium situated on the pseudopodium which is 
still surrounded by the perichaetium ; to the left is a barren branch. F Portion of a 
foliage-leaf seen from above: I perforations; b chlorophyll-containing cells; s spiral 

colourless. This anatomical structure is an essential condition for 
+be formation of peat. The Bog-Mosses grow by preference on 


moors, which they cover with a thick carpet saturated with water. 
The lower extremities of the plants perish very rapidly, and 
gradually become converted into peat, and the branches thus 
separated from each other become independent plants. The 
sporangia (Fig. 199 D, E) are spherical, but with a very short 
stalk. They open by a lid, but have no annulus. The archegonium 
(Fig. 199 C) persists at the base of the sporogonium as in the 
Liverworts. Only one genus, Sphagnum. 

Family 2. Schizocarpese. 

The Mosses which constitute this family are of a brownish-black colour and 
are found living on rocks. The sporangium resembles that of the Liverworts 
inasmuch as it opens by four valves, but these continue attached to each other at 
the apex as well as at the base (Fig. 193). There is only one genus : Andreata. 

B. The stalk is formed from the lower portion of the sporogonium. The 
columella is continued to the summit of the sporangium and united with it 
(Archidium has no columella.) 

Family 3. Cleistocarpeae. 

The fruit does not dehisce in the regular way, but the spores are liberated by 
decay. They are small Mosses which remain in connection with their protonema 
until the sporangium is mature. The archegouium remains sessile at the base 
of the short capsule-stalk, and is not raised into the air (compare Hepaticae). 
Phascum, Ephemerum, Archidium, Pleuridium. 

Family 4. Stegocarpese. 

To this belong the majority of the Mosses, about 3,000 species. 

The capsule opens as in Sphagnum by means of a lid (operculum) , 
which is often prolonged into a beak. Round the mouth of the 
opened capsule, a number of peculiar yellow or red teeth are to be 
found. These constitute the peristome ; their number is four, or a 
multiple of four (8, 16, 32 or 64). The form and thickenings 
of these teeth are widely different, and on this account are used by 
Systematists for the purposes of classification. In some Mosses 
(Fig. 200 C, D) there is a double row of teeth. Except in 
Tetraphis they are not formed from entire cells, but from the 
strongly thickened portions of the wall of certain layers of cells 
belonging to the lid, and persist when this falls off. They are 
.strongly hygroscopic, and assist greatly in the ejection of the lid, 
in which operation they are considerably aided by a ring of elastic 
cells with thickened walls, situated in the wall of the lid near the 
base of the teeth. This ring is known as the annulus. The arche- 
gonium is raised into the air like a hood, the calyptra, which 


either covers the sporangium on all sides (having the shape of a 
bell), or is split on one side (Fig. 200 jB, 7i). 

Among peculiar forms may be mentioned : Splachnum, which is especially 
remarkable for the collar-like expansion at the base of the capsule. Fissidens 

deviates in having a flat stern and leaves 
arranged in two rows. The leaves are 

I boat-shaped and half embrace the stem. 

Schistoatega has two kinds of sterns. 
The barren ones resemble Fern-leaves ; 
they have two rows of leaves, which are 
attached together vertically, are decur- 
rent, and coalesce at their bases. The 
fertile ones have an ordinary appearance. 
Tetraphis : the peristome is composed 
of four teeth, wbich are formed from entire 
cells. 2'. pellucida has peculiar gemmae. 

The family is divided into two 
groups : the Musci acrocarpi, the 
growth of whose main axis is 
limited and terminated by the 
formation of the sexual organs ; 
and the Musoi pleurocarpi, whose 
sporogonia are situated on special 
lateral shoots, while the growth of 
the main axis is unlimited. 

FIG. 200. A Hypnum populeum. B 
and C Sporangia, with hood (h), and 
operculum (!'), and without these (C), 
showing the peristome (p). D The 
mouth of the capsule of Fonfinalis ant-i- 

A. Acrocarpi. 

Order 1. Weisiaceae. Peristome, 
with 16 teeth arranged in one series, rarely 
wanting. Leaf with midrib. Campy lopus, 
Dicranum (D. scoparium, common in forests), Dicranella, Cynodontium. 
Weisia, Gymnostomum (no peristome), Systegium. 

Order 2. Leucobryaceae. Peristome with 16 teeth. Leaves with three or 
more layers of cells, of which the external ones are air-conducting and per- 
forated (as in the Sphagneas), the middle one containing chlorophyll. Leuco- 

Order 3. Fissidentaceas. Peristome as in the preceding ones. The 
leaves are arranged in two rows on the plagiotropic shoots ; in Fissidens the 
midrib of the leaf bears wing-shaped outgrowths. Conomitrium, Fissidens. 

Order 4. Seligeriaceaa. Peristome with 16 undivided teeth. Very small 
Hock-mosses. Seligeria. Blindia. 

Order 5. Pottiacese. Peristome with 16 teeth, which are divided almost 
to the base, or with 32 teeth. Calyptra hood-like. Barbula (B. muralis, B. 
raralis), Trichostomum, Leptotrichiiin. Ceratodon pnrpureus. Distichium. 

MUSCINE^. 197 

Order 6. Grimmiaceae. The leaf-cells are often papillose ; in the upper 
portion of the leaf, small, and of roundish shape. The calyptra is most 
frequently hood-like or conical. Eucalypta. Orthotrichum, often with short- 
stalked capsule, is found on trees. Coscinodon.Hedwigia.Grimmia, Raconii- 
trium. Cinclidotus. 

Order 7. Schistostegaceae. The stems are of two kinds (see above) ; 
Schistostega osmundacea, in caves, has a bright emerald protonema. 

Order 8. Splachnaceee. The capsule has a large, collar-like neck (see 
above). Splachnum (especially on manure). 

Order 9. Funariaceae. Capsule pear-shaped. Ftmaria (F. hygrometrica has 
a very hygroscopic seta, becoming twisted when dry, and straightening with 
moisture) ; Physcomitrium ; Discelium. 

Order 10. Bryaceae. The capsule is thicker towards the apex; most 
frequently pendulous. Philonotis, Bartramia. Aulacomnium. Paludella 
Meesea. Mnium. Bnjum, Webera, Leptobryum. 

Order 11. Polytrichaceae. Single peristome, formed by 16, 32, or 64 teeth. 
Leaves with longitudinal lamellae on upper surface. Polytrichum has long, 
hairy calyptra. Catharinca (C. undulata, in forests). 

Order 12. Georgiaceae. Peristome with 4 teeth (see above). Tetraphis 
(T. pellucida has gemmae). 

Order 13. Buxbaumiaceae. Capsule asymmetrical; double peristome: 
the interior one conical, with 16 or 32 longitudinal folds. Buxbaumia (B. 
aphylla) ; Diphyscium. 

B. Pleurocarpi. 

Order 14. Fontinalaceae. Long, floating Water-Mosses. Fontinalis (F. 
antipyretica is found in streams). Dichelyma. 

Order 15. Hookeriaceae. Pterygophyllum. 

Order 16. Leskeaceae. Dull-looking Mosses, with papillose or warted 
leaves. -Thuidium, T/tuj<i-like with regularly arranged 1-3 doubly pinnate 
stems ; Anomodon, Leskea. 

Order 17. Pterogoniaceae. Pterigynandrum filiforme, etc. 

Order 18. Fabroniaceae. Anacamptodon. 

Order 19. Neckeraceae. Stems most frequently with flat, leafy branches. 
The leaves are smooth, never with longitudinal folds. Neckera. 

Order 20. Hypnaceae. The leaves are smooth with square, often bladder- 
like, cells at the edge. Hylocomium (H. splendeiis, H. triquetrum) ; Hypnum ; 
Brachythecium Plagiothecium. Eurhyjtchium.Homalothecium, Isothecium, 
Orthothecium, Homalia. Climacium, Lescurcea, Leiicodon. 

The Mosses occur all over the globe. Many are found in great numbers, and 
growing thickly massed together, they form an important feature in landscapes 
(for example Sphagnum and Polytrichum in the Arctic Tundra). In the Northern 
and Arctic regions the Mosses are very plentiful, and often form a considerable 
part of the vegetation, while in the Tropics they are insignificant. 

Species of Hypnum and Polytrichum, like Sphagnum, play an important part 
in the formation of peat. 



The alternation of generations is as distinct in this Division as 
in the Mosses, but the sexual generation consists of only a small 
thallus, the prothallium, which bears directly the sexual organs, 
antheridia and archegonia ; and the asexual generation, which 
arises from the fertilisation of the oosphere, is no longer a single 
short-lived sporangium, but a highly developed, generally per- 
ennial, plant provided with stem, leaves and true roots (Ferns, 
Horsetails, etc.), the sporangia being borne on the leaves. In this 
latter generation the tissues are differentiated into epidermis, 
ground tissue and vascular tissue; in the last named the bundles 
are closed, and in the majority of cases concentric. 

The sexual generation, gametophyte, or prothallium, is 
always a thallns, although not always green and leaf-like (Figs. 

205, 215, 222, 229, 235, etc.) It is very small, even in cases where 
it attains the greatest development, and consists only of paren- 
chymatous cells. The prothallium is nourished by hair-like roots 
(rhizoids) and has only a transitory existence, dying soon after the 
fertilisation of its oosphere. 

The ANTHERIDIA exhibit great variations in structure which, 
however, must be considered as modifications of the fundamental 
type which is found in the Mosses. These modifications will be 
mentioned under the various families. The spermatozoids are 
always spirally-coiled, self-motile, protoplasmic bodies, with most 
frequently a large number of fine cilia on the anterior end (Figs. 

206, 223, 234). They are formed principally from the nucleus of the 
mother-cell, and portions of the cytoplasm often remain for a 
time attached to their posterior end. 

The ARCHEGONIA are more uniform throughout the entire 
Division, and more closely resemble those of the Mosses. They 
are, as in the previous Division, principally flask-shaped ; but the 




central portion, which encloses the oosphere, is always embedded 
in the tissue of the pro thallium, so that the neck, which is formed 
of 4 rows of cells, projects above the surface (Figs. 201 3 , 222 //). 
The development of the archegonium in a Fern is seen in the 
accompanying figure (Fig. 201). The archegoniumis developed from 
a surface cell, which divides into three cells by two walls in 
a direction parallel to the surface of the prothallium (Fig. 201). 
The most internal cell becomes the ventral portion of the arche- 
gonium. The external one (fr) divides perpendicularly to the sur- 
face of the prothallium into four cells, which again divide parallel 
to the surface and form the neck (b, in 2 and 3). The interme- 
diate cell projects upwards into the neck and divides into two, 

FIG. 201. Pteris serrulata. Development of archegonia. 

the lower one, after the separation of the ventral canal-cell, becom- 
ing the oosphere, arid the upper one the neck-canal-cell (c, in 2 and 3). 
As in the Mosses, the divisional walls of the neck-canal-cells 
become mucilaginous, causing the rupture of the neck of the 
archegonium. Fertilisation takes place as in the Mosses, and the 
passage of the sperm atozoids, along the neck, to the oosphere, 
has been observed. Water (rain or dew) is similarly necessary for 
the movements of the spermatozoids, and hence for fertilisation. 
The other classes of the Division chiefly deviate from the Ferns 
in having the archegonium sunk deeper into the prothallium, and 
the neck reduced in length (compare Fig. 201 with Figs. 216, 222, 
235, 236). 


According to the nature of the spores, the three classes of 
the Vascular Cryptogams are each divided into isosporous and 
heterosporous groups. 

I. The isosporous Vascular Cryptogams hare only one kind 
of spore. The prothallium developed from this is in some cases 
monoecious, bearing both antheridia and archegonia; but in others 
there is a distinct tendency for each prothallium to bear only 
antheridia or archegonia (dioecious) true Ferns and Lycopodium. 

In Equisetum there is only one kind of spore, but two kinds of 
prothallia are developed, one of which bears only antheridia 
(male), the other only archegonia (female) ; but the one that 
bears antheridia may be transformed into the one that bears 
urchegonia and vice versa. 

II. In the higher group, heterosporous Vascular Cryptogams 
(Selaginella and Isoetes, etc.), there are two distinct kinds of 
spores, the small, microspores, and the large, macrospores. The 
microspores are male, and produce prothallia which bear only 
antheridia. The macrospores are female, and produce prothallia 
which bear only archegonia. 

Corresponding to this difference in the spores, there is also 
found a difference in the development of the prothallium. In the 
Isosporea? the prothallium is large, and either green, leaf-like, and 
provided with rhizoids (most of the Ferns, Horsetails, etc.), or 
subterranean, pale-coloured, and globular (Ophioglossum, Lycopo- 
dium). It lives vegetatively for a fairly long time, and generally 
produces a large and varying number of archegonia and an- 
theridia. The prothallium in the Heterosporese is gradually more 
and more reduced, its independent and vegetative life becomes of 
less and less importance, it becomes more dependent on the rnother- 
plant, and projects from the spore very slightly, or not at all. 
The antheridia and archegonia become reduced in number to one, 
and also degenerate in point of development. 

It may hei-e be remarked that the gradual development of the 
asexual generation, the development of the two kinds of spores, 
and the progressive reduction of the prothallium and sexual 
organs which is found in this Division, is continued to the Gymno- 
sperms and Angiosperms. The microspores are in these called 
pollen-grains, and the male prothallium is very rudimentary. 
The macrospores are termed embryo-sacs, and the female prothal- 
lium, the endosperm. 

The asexual generation, sporophyte. When the oosphere, 


which in this case as in all others is a primordial cell, is fer- 
tilised, it surrounds itself with a cell-wall and commences to divide 
into a number of cells, to form the embryo. 

The first dividing wall (basal wall) is nearly horizontal, and in the direction of 
the longitudinal axis of the archegonium. The next wall is vertical, and the 
next perpendicular to the other two. The oosphere, therefore, is now divided 
into eight octants by these three walls. The basal wall divides the embryo into 
a hypobasal and an epibasal half. From the first one, by continued divisions, 
the first root is developed ; from the latter, the stem and leaves. After the for- 
mation of the octants the development proceeds in somewhat different ways in 
the various classes. In addition to the stem, leaf, and root, a " foot " is deve- 
loped from the hypobasal half which remains enclosed in the prothallium, and 
conveys nourishment from the prothallium to the young plant until it is able 
to sustain itself (Fig. 202). The formation of these members in the embryo 
depends on the position of the oosphere in the archegonium and prothallium, 
and is independent of gravity. 

FIG. 202. A&iantum capillas veneris. Vertical section through a prothallium (//), 
with a young plant attached on its under side (mag. about 10 times) ; r the first root, and 
b the first leaf of the young Fern-plant ; m the foot. In the angle between m. and & lies the 
apex of the stem : Ii the rhizoids of the prothallium ; ae ce unfertilised archegonia. 

In the Mosses the asexual generation is the sporogonium, which 
is limited in its development and in a great measure dependent 
upon the sexual generation, upon which it is situated ; but in the 
Pteridophyta this generation is an independent and highly developed 
plant, provided with stem, leaf, and true roots, and has in many 
instances an unlimited development. The Pteridophyta are the 
lowest Division with true roots. The root which is first formed is 
very similar in nature to the primary root of the Monocotyledons ; 
it very soon dies and is replaced by others which are more per- 
manent, and developed upon the stem (adventitious roots) ; roots 
are wanting in Saluinia, Psilotum, and some Hymenophyllacese. 
The differentiation is, however, not so complete as in the Flower- 
ing-plants, and so many leafy forms are not found. The various 
members of these plants are anatomically much higher than in the 



Mosses, having an epidermis, a ground tissue with variously dif- 
ferentiated cells, and a highly developed vascular system. The 
vascular bundles, like those in the Monocotyledons, are without 
cambium, and closed ; they are therefore incapable of any increase 
in thickness. In general the bundles are concentric, with the 
bast round the wood (Fig. 203). The wood is almost entirely 
made up of scalariform tracheides. 

In Iso'ftes a secondary thickening takes place by a cambium, which is formed 
inside the cortex, constructing secondary cortex to the exterior, and secondary 
wood towards the interior. Botrychium has also a thickening growth. Collateral 
vascular bundles occur in Osmundacecc, Equisetacece, and the leaves of many 
Pulypodiacece, etc. 

It is a point of special interest, that the gigantic forms of Ferns, 

Equisetums, and Club- 
Mosses (which flourished 
in earlier geological 
periods, when these classes 
attained their highest de- 
velopment) possessed 
some means of increasing 
in thickness. 

The sporangia are in 
all cases capsule-like, and 
burst open when ripe to 
eject the spores. They 
are nearly always situated 
on the leaves (in Li/copo- 
diacece, in the axils of the 
leaves, or above these, on 
the stems themselves). In 
some forms (LEPTOSPOR- 
ANGIATJE), the sporangia 
are developed from a single epidermal cell ; in others (EusPOR- 
ANGIATVE), from a group of epidermal cells, or from cells which 
lie beneath the epidermis. In the first group a primitive mother- 
cell (archesporium) is formed, which divides commonly into sixteen 
special mother-cells. In the latter group, on the other hand, a 
number of primitive spore-mother-cells are developed. In each 
sporangium three different tissues are generally developed ; an 
innermost sporogenous one (s in Fig. 20 i A), which arises from 
the archesporangium ; an outermost one, which forms the ivall (a), 

FIG. 203. Portion of the stem of a Fern. Above 
is seen the transverse section, with vascular bundles 
of different form and size. The rhombic figures on 
the side of the stem are leaf -scars. 



and may be one or, more rarely, several layers in thickness ; and 
an intermediate one, the tapetum (Fig. 204 A, B, b ), which is rich 
in protoplasm, and Avhose cells are dissolved so that the spores 
float freely in the fluid thus provided. The spores arise as in the 
Mosses (in tetrads), by the cross-division of the special mother- 
cells, and according to the manner in which they are arranged 
in the mother-cell have either a tetrahedral form, with a large 
base resembling a segment of a ball, or are oblong (bilateral 
spores). Their construction is the same as in the Mosses (p. 

The spore-formation in its earliest commencement takes place in 
the same way in the Isosporous and the Heterosporous Vascular 

Fr&. 204. Selaginella in&qualifolia. A A yonng sporangium, which may develope either 
into a macro-, or a micro-sporangium. B A microsporangium. 

Cryptogams; but from a certain point, after the tetrahedral divi- 
sion, a difference occurs with regard to the macrosporangia. All 
the spores formed in the microsporangium may complete their 
development ; but those which are formed in the macrosporangium 
are generally aborted, with the exception of one or four, and these 
consequently attain a much larger size (see Fig. 239. The series to 
the left are microsporangia ; those to the right, macrosporangia). 

APOGAMY. In some Ferns (Pteris cretica ; Aspidiitm filix mas, var. cristatum; 
A. falcatum; Todea africana) the young plant is not developed as a consequence 
of fertilisation, but as a bud from the protballium. This is known as apogamy, 
or loss of the power of sexual reproduction. The antheridia are generally 
more or less developed ; archegonia are entirely wanting in Asp. filix max, var. 


cristatum. This variety has probably only become apogamous through cultiva- 
tion. Many specimens of Isoetes lacustris, in a lake in the Vosges mountains, 
produce in the place where the sporangia are usually found, a vegetative shoot 
which grows into a new plant, so that the sexual generation is wanting in 
this case. Some specimens have sporangia on some leaves, and shoots on 

Apospory, or the formation of prothallia instead of sporangia and spores on 
the leaves, is found in Athyrlum filix femina, var. clarissimum. In this case 
the development of the sporangia proceeds only to a certain point, and from 
these arrested sporangia the prothallia are produced. Normal sporangia are 
entirely wanting in this variety, and in Aspidium angular?, var. pulcherrimum, 
sporangia are completely wanting. Compare the Mosses (page 188). 

The Vascular Cryptogams are divided into three large classes, 
in each of which a progressive development can be traced from the 
isosporous to the heterosporous forms, but some of these are now 
only known as fossils. 

Class 1. Filicinae (Ferns). The stem is small in comparison 
with the leaves, and branches only seldom, and then by lateral 
shoots. The leaves are scattered, large, often deeply divided, and 
of various highly developed forms. The undeveloped leaves are 
rolled up in the bud, having what is termed circinate venation. 
The sporangia are situated on the edge or on the lower side of 
the leaves, those on which the sporangia are borne (sporopliylls} 
being often the ordinary foliage-leaves ; but in a few cases the 
fertile differ from the barren ones (a higher stage in development). 
The fertile leaves are not confined to definite parts of the shoot, 
and do not limit its growth. The archesporium is most frequently 

A. Isosporous: Sub-Class 1. Filices (True Ferns). 

B. Heterosporous : Sub-Class 2. Hydropteridea? (Water 

Class 2. Equisetinae (Horsetails), in its widest meaning. 
The leaves in this class are small in comparison with the stem. 
They are arranged in whorls, and unite to form a sheath. The 
sporangia are situated on specially modified, shield-like leaves, 
which are closely packed together and form a " cone." The cone 
is borne terminally, and limits the growth of the shoot. The 
sporangia are developed from a large group of epidermal cells, the 
archesporium being unicellular. The branches are arranged in 
whorls, and develope acropetally. 

A. Isosporous: Sub-Class 1. Equisetaceoe. Existing forms. 

B. Heterosporous : Sub-Class 2. Extinct forms. 


Class 3. Lycopodinae (Club-Mosses). --Roots generally 
branching dichotomously. The leaves are scattered or opposite, 
and in proportion to the stem very small, undivided, and simple. 
They are scale-like and triangular, tapering from a broad base to 
a point. The sporangia are situated singly (except in Psilotacece), 
and almost in every case on the upper side of the leaf or in the 
axil of a leaf; but in some cases they are borne on the stem, just 
above the leaf-axil. The sporangia arise from groups of epidermal 
cells. The sporophylls are often modified, and differ from the 
foliage-leaves ; they are then arranged in cones placed terminally 
on branches, thus limiting their growth. 

A. Isosporpus : Sub-Class 1. Lycopodieas. 

I?. Heterosporous : Sub-Class 2. Selaginellese. 

Class 1. Filicinae (Ferns). 

The characteristics of this class have already been given on page 

The class is divided into two sub-classes : 

1. The TRUE FERNS, FILICES, have one kind of spore which 
generally developes monoecious prothallia, relatively large and 
green. The sporangia are most frequently situated in groups 
(sori), which are often covered but not enclosed by an indusium. 

2. WATER FERNS, HYDROPTERIDE.E, have micros porangia with 
many (4x16) microspores, and macrosporangia, each with one mac- 
rospore. The prothallium is small, and projects but slightly from 
the germinating spore. The sporangia are situated in groups 
(sori), which are either enclosed by an indusium, or enveloped in 
a portion of a leaf, to form " fruits " termed sporocarps. 

The old name for the Hydropterideas, "Rhizocarpeaa," i.e. the " root-frnited," 
originated from the erroneous supposition that the sporocarps were borne on 
the roots. 

Sub-Class 1. Filices (the True Ferns). 

Of the eight orders (with about 4,000 species) comprised in this 
sub-class, the Polypodiaceas is the largest (having about 2,800 
species) and the most familiar; for this reason it will be taken as 

The sexual generation. When the spore germinates, the 
external covering (exospore) is ruptured, as in the Mosses. The 
internal cell-wall (endospore) grows out as a filament, which soon 
divides and gives rise to the prothallium, a flat, cellular expansion 
resembling the thallus of a Liverwort. In its fully developed state 



the prothallium is generally heart-shaped, dark green, and pro- 
vided with root-hairs, and it attains a 
diameter of about one centimetre (Fig. 
205). It is formed of one layer of cells, 
except along the central line near the 
anterior depression, where it becomes 
several layers of cells in thickness, 
forming the " cushion," on the lower side 
of which the archegonia are developed. 
The antheridia are first formed ; they 
are thus found 011 the oldest parts of the 
prothallium, on its edge, or among the 
root-hairs. The archegonia are developed 
later, and are therefore found near the 
apex. Several tropical Ferns have pro- 
thallia 1 deviating from this typical form; 
Trichomanes (Order Hymenophijllace^) 
has filamentous, branched prothallia, 
which resemble the protonema of a Moss. 

Others, again, have strap-shaped prothallia, which resemble the 

thallus of certain Liverworts. 

FIG. 205. Prothallium (j>p) of 
Maiden hair (Adiantum capllliis 
venevis) with a young plant 
attached: bfirstleaf; w' primary 
root; w" adventitious roots; h h 
root-hairs of the prothallium 
(x abt. 30). 

Fio. 206. Antheridia of Maiden-hair ( x 550) . A Unripe ; B ripe, but unopened ; C open 
and ejecting the sperrnatozoids (s). Those which have been last ejected are still lying en 
closed in their mother-cells, the others are coiled up and drag with them the c> toplasmic 
remains (b) ; /cells of the prothallium. 

1 In the Polypodiaceas unisexual prothallia as distinct as those of Equisctum 
are of common occurrence. 


The ARCHEGONIA have been already mentioned (p. 199, Fig. 201). 
The ANTHERIDIA are hemispherical or slightly conical bodies (Fig. 
206). They consist, as in the Mosses, of a wall formed by one 
layer of cells, which encloses a number of spermatozoid-mother- 
cells (A and B). The antheridia when ripe absorb water, and 
are ruptured, and the spirally-coiled spermatozoids liberated (Fig. 
206 S). The spermatozoids have been observed to pass down the 
neck of the archegonium, and to fuse with the oosphere. 

The asexual generation. The first leaf, the "cotyledon," 
of the embryo developed from the oospore (Figs. 202, 205) is 
always small, and has a very simple shape. The leaves which 
occur later become more perfect, stage by stage, until the perma- 
nent form of leaf has been attained. The STEM is most frequently 
a subterranean or a semi-aerial rhizome ; it is only in the tropical, 
palm-like Tree-Ferns, that the stem raises itself high in the air and 
resembles that of a tree, with leaf-scars or with the remains of 
leaves attached (Figs. 207, 203) ; in certain species the stem is en- 
cased in a thick mat of aerial roots (Dicksonia antarctica). When 
the rhizome is horizontal the internodes are frequently elongated, 
and the leaves are arranged in two rows, as in Poly podium vulgare 
and in the Bracken-Fern (Pteridium aquilinum), etc. ; it is also 
generally dorsiventral, having a dorsal side on which the leaves are 
situated, and a ventral side, different from the former, on which 
the roots are borne. When the stem ascends iu an oblique direc- 
tion, or is nearly vertical, its internodes are extremely short, and 
the leaves are arranged in a spiral line with a complicated phyllo- 
taxis, e.g. in Atliyrium filix-fceniina, Asyidium jilix-mas, etc. The 
BRANCHING upon the whole is extremely slight, and is generally 
confined to the petiole (e.g. Aspid. filix-mas), or to the stem near 
the insertion of the leaves. Several species normally form buds on 
different parts of the lamina. The buds which are formed on the 
stem are not confined to the leaf -axil as in the higher plants. 
The Tree-Ferns, generally, do not branch at all. 

The VASCULAR BUNDLES are concentric, with the wood surrounded 
by the soft bast. In tranverse section they are seen as circles or 
irregularly-shaped figures (Fig. 203), the name of " King Charles 
and the Oak" (Bracken-Fern) having originated from the appear- 
ance which the bundles present in oblique section. In Osmunda 
they are collateral and resemble those of the Flowering-plants. 
Round each individual bundle is often a sheath of thick- walled, 
hard, brown, sclerenchymatous cells, which act as a mechanical 



tissue; similar strands are also found in other parts of the 


The LEAVES in nearly all species are only foliage-leaves, borne 

FIG. 207. Various Ferns (1, 2, 3, 4). 

in a spiral. They have an apical growth which continues for a 
long time, and some require several years for their complete de- 
velopment. In the buds they are rolled up (circinate) ; not only 
the midrib, but also all the lateral veins, and even the terminal 


portions of a leaf are sometimes rolled up together, the tissues of 
the leaf being already folly developed and only waiting to expand. 
The leaves are often excessively divided and compound, with 
pinnate branches, and have an epidermis with stomata and a well- 
developed system of venation. Stipules are only found in Marat- 
tiacece and OphioylossaceiP. 

Very often peculiar hairs or scales (pale<z, ramenta), dry, brown, 
flab and broad, are found on stem and leaf. 

The SPORANGIA are small, round capsules, which, in a very large 
number of Ferns, are formed on the back, but more rarely on the 
edge of the ordinary foliage-leaves. It is very seldom that there 
is any difference in form between the barren foliage-leaves and the 
fertile leaves, as is found for example in Blechnnm spicant or 
Struthiopteris ; or that the fertile part of the leaf is differently 
constructed from the barren portion of the same leaf, as in the 
Royal-Fern (Osmunda). In such instances the mesophyll of the 
fertile parts is poorly developed. 

The sporangia in the Poli/podiacece are lens-shaped, with long 
stalk (Fig. 211 1}) : their wall consists of one cell-layer on which 
a single row of cells, passing vertically over the top (that is along 
the edge of the sporangium), is developed into the "ring" (annu- 
lus). The cells of the annulns are very much thickened on the 
inner and side walls, and are yellowish-brown. The thickened 
cells, however, do not entirely encircle the sporangium, and on 
one side, near the stalk, they pass over into large, flat, thin-walled 
ells. These form a weak point in the wall, and it is here that 
the sporangium is opened diagonally by the elongation of the 
annulus. The sporangium of the Polypodiacea? opens as it dries. 
The cells of the annulus are very hygroscopic, and in straighten- 
ing, the annulus bends back with a jerk, thus ejecting the spores 
to considerable distances. The cells of the annulus absorb water 
with great readiness. [The sporangium arises as a single epi- 
dermal cell, from which a basal stalk-cell is cut off. Three oblique 
cell-walls, intersecting near the base, are next formed in the upper 
cell, and a fourth between these and parallel to the free surface ; 
an inner tetrahedral cell enclosed by four others is thus formed, 
the outer cells become the wall of the sporangium, while the inner 
cell, by a series of walls, parallel to its sides, cuts off a layer of 
cells which eventually form the tapetum, the remaining central 
cell constituting the archesporium.] 

The SPORES are either oblong and bilateral, or they are tetra- 

w. B. P 


liednc. with curved sides, depending upon the way in which the 
tetrad division has taken place. 

The sporangia are almost always situated on the nerves and 
gathered into groups, sori, which differ in form in the various 
genera. The sori, in many genera, may be covered by a scale- like 
structure, the indusium (Figs. 211 B, 212). 

In the majority of cases, each sorus is situated on a small 
papilla (placenta, or receptacle), which is supplied by a small vascu- 
lar bundle. Between the sporangia, hairs (paraphyses) are often 
situated, which spring either from the placenta or from the stalks 
of the sporangia. 

Systematic Division. The Ferns may be divided into two 
groups, characterized by the structure and development of the. 
sporangia. The sporangia in the EUSPORAMUAT.E take their origin 
from a group of epidermal cells, and their walls are formed by 
several layers of cells. The archesporium is the (not tetrahedric) 
hypodermal terminal cell of the axial row of cells which give rise 
to the sporangium. In the LEPTOSPORANGIAT2E the sporangia are 
developed from single epidermal cells, and their walls are uni- 
layered. The archesporium is a central, often tetrahedric cell, 
from which sixteen spore-mother-cells are developed. 1 It is diffi- 
cult to say which form is the oldest (according to Prantl, those 
which have the sori on the nerve-endings) ; however, the Eusporan- 
giatse would seem to have made their appearance long before the 
others, and also well defined Marattiacese and Ophioglossaceos 
occur in the Kulm and Coal period, before the true Polypodiacese. 

About 4,000 species of Ferns are now existing, and they are 
found especially in tropical and sub-tropical forests. 

Family 1. Eusporangiatae. 

Order 1. Ophioglossaceae. The prothallium differs from 
that of all other Ferns in being subterranean, free from chlorophyll, 
pale and tuberous. The stem is extremely short, with short 
internodes, most frequently unbranched, vertical, and entirely 
buried in the ground (Fig. 208 s/). In several species (among 
which are the native ones) one leaf is produced every year, which 
lias taken three to four years for its development. In Botry- 
rliinm a closed, sheath-like basal part of each leaf covers the sub- 
sri|iit'iit leaves during their development. In Ophioglossum and 

The position of the annulus varies in the different orders ; longitudinal 
in I'olypodiacere, Hymenophyllaceae, and Cyatheacea- ; transverse iu Schizaj- 
aceiv, (ilrichrman ;r : indistinct or apical 'in Osmundaceae, Ophioglossaceae, 
Marattiaceee, Sulviniaci-jf, Marsiliaceae. 



others each leaf has at its base an intrapetiolar, cap-like sheath, 
which protects the succeeding leaf. The leaves are of two kinds : 
(a) foliage, which in Ophioglossum vulgatum are lanceolate and 
entire, but in Botrychinm however, are pinnate (b in Fig. 208 
A, B} ; and (6) fertile, which are found facing the upper side of 

FIG. 209. - 
Fertile leaf of 

FIG. 208. .4 Ophioglossum vulgatum (Ailfler's-tongue) : B Botryeliium lunar/a (Moon- 
won), both natural size ; r-r roots ; bs leaf -stalk ; st stem ; b foliage-leaf ; /fertile leaf. 

the foliage-leaves. These latter in Ophioglossum are undivided 
and spike-like (Fig. 209 A), but pinnate in Botrychium (Fig. 208 B). 
Each foliage and fertile leaf are branches from the same petiole. 
The large sporangia are placed laterally, and open by two valves. 
No annulus is formed (Fig. 209). Ophioglossum reproduces vege- 
tatively by adventitious buds on the roots. 



Three genera with about twelve species. 

Order 2. Marattiacese are tropical Ferns, whose gigantic 
leaves resemble those of the Polypodiacese, but have stipules in 
addition. The sporangia are grouped in sori, situated on the lower- 
side of the leaves, the sporangia in each sorus being arranged 
either in two rows or in a ring. In Angiopteris they are isolated 
(Fig. 210 A), but in the other species (Kaulfussia, Dauct>a, Marattia\ 
they are united, and form " synangia " divided into a number of 
chamber's corresponding to the sporangia. These open by clefts 
or pores. Marattin presents the highest development, as its 
sporangia are completely united in a capsnle-like synangium, 
which is closed until maturity, and then opens by two valves. 
In each valve there is a row of three to eleven sporangia, each 
opening by a slit towards the inside (Fig. 210 B, C). An indusium 
encloses the sorus, except in Kanlfussia ; it is formed of flat and 
lobed hairs, which resemble the hairs of the other portions of the 

leaves. In Angiopteris and Mar- 
attia the indusium is very rudi- 
mentary ; in Dancea it forms a 
kind of cupule. 

The numerous fossil Marattiacete 
(15 genera, with 98 species) present 
similar differences to those now living, 
but more various forms are found, for 
example, with solitary free sporangia. 
Those now living are the last small 
remnant (4 genera with only 23 
species) of a once dominant family, 
which existed from very early times, 
and whose culminating point was 
reached in the Kulm and Coal periods. 

The Ophioglossaceas appear also in 
the Kulm and Coal periods, and were 

FIG .210. Sporangia of the Marattiaceae : 
.4 Angiopteris; B and C Marattia ; C is a 
half sorus with nine sporangia, each of 
which has opened by a longitudinal cleft. 

about as numerous as at the present time (presumably 2 genera, with 19 
species). Leptosporangiate Ferns appear however to have occurred first of all 
in the Trias-formation. 

Family 2. Leptosporangiatae. 

Order 1. Polypodiaceae. Sporangia on the lower side of the 
leaves, stalked and provided with a vertical, incomplete annul us ; 
dehiscing by a transverse cleft (Fig. 211 !>). The genera are 
distinguished by the form of the indusium and the position of 
the sori, etc. 



1. The sporangia cover the entire lower surface of the leaf 
(Tropical America and Asia). Acrostichum, Platycerium. 

2. Sori without indusia, circular or oval. Polypodium (Fig. 211 
A). The leaves are most frequently situated in two rows on the 
dorsal side of the creeping rhizome, and fall off leaving a smooth 
scar behind. P. vulgare, common in woods, on stones. (Pliego- 
pteris also has no indusium ; see page 214). 

3. The sporangia are situated in continuous lines just inside the 
margin of the leaf. Pteris 1 : the sporangia form a continuous line 
along the entire margin of the leaf (Fig. 211 C), which bends over 
and covers the sporangia, forming a " false-indusium." Pteridium 
has linear sori situated on a marginal vascular bundle, covered 
by two linear basal indusia, of which the outer is bent over like 
the edge of a leaf. P. aquiliuum (Bracken,) has a wide-spreading 

PIG. 211. Portions of leaves with sori. A Polypodium. B Aspidium. C Pteridium. D 
A sporangium of one of the Polypodiacea? : r the annulus ; s spores. 

rhizome with large alternate leaves, placed on opposite sides, at 
some distance apart. Only one leaf is developed from each branch 
every year. 

Adiantum (Maiden-hair) : sori on the underside of small portions of the edge 
of the leaf, which are beat over (false indusium). Cnjptogramme (Allosorus), 

4. The sori are oval or linear, situated on one side of the 
vascular bundle. Asplenium (Fig. 212 A) . sori linear; indusium 
with one of its edges attached at the external side. A. ruta mura- 
ria (Wall-Rue) ; A. septentrionale; A.trichomanes. Athyrium: sori 
linear cr curved; A. jilix-fwmina (Lady-Fern). Scolopendrium 
1 The former genus Pteris is divided into Pteris and PteriiUiiin. 



(Fig. 212 if) : sori as in Asplenium, but situated in pairs across 
the lanceolate, entire leaves. Each sorus is covered on the ex- 
ternal side by an indusium, whose free edges are parallel and ap- 
proach each other. S vulijare (Hart's-tongue). BlecUmim (B. tpicant, 
Hard Fern ; the fertile leaves differ from the barren, the pinna? being narrower, 
while tbe underside is almost entire'y covered with sori, arid hence they are of 

a much darker brownish hue than the barren ones) Ceterach : iudusium 

rudimentary or absent. 

5. Sori circular and covered by a shield-like, or reniform 
indusium. Aspidium (Fig. 211 B) ; the leaves wither away and 
leave no scar upon the root-stock. A. Jilix-mas (Male-Fern) ; A. 
tt^inulosum. Phegopteris has no indusium, the withered bases of 
the leaf-stalks are persistent ; P. dryopteris and P. polypodioides. 

6. The indusium is situated below the sori, and has the shape of 
a one-sided scale (Cystopteris, Struthiopteris), or of a cup or cupule, 
which in Woodsia is sometimes fimbriate (Fig. 212 C, !>}. 


FIG. 212. A Asplenium. B Scolopendrium. C Woodsia ; D single sorus of the same, 
/v ( 'i/at/icu : the sporangia have fallen off in the upper sori. (All magnified.) 

7. The sori are situated on the margin of the leaf, and at the end of a 
vascular bundle. Indusium, semi-cupjlar. Davallia. Principally tropical 
species. 1 in S. Europe. 

This order is the greatest, comprising about 2,800 species, the 
majority being perennial plants. A few are large, and known as 

As plants in conservatories and rooms the following are cultivated : species of 
<;i/>nmjt (inline (tropical America), Lomaria, Nephrolepis, Pteris (P. scrrulata, 

Officinal. Aspidium Jilix-mas, rhizome and the withered petioles. Species of 
Alsophila and Cibotiuin give Penghawar Djambi. The rhizome of Pteridium 
aquilinum, var. esculentuin, contains so much starch that it is used as food. 

The other orders of true Ferns deviate from the Polypodiaceae, especially in 



the formation of the annulus, the bursting of the sporangium and its mode of 
attachment and development, and in the differences in the formation of the pro- 
thallium, etc. The principal are : 

Order 2. Hymenophyllaceae. To this order belong the lowest and most 
Moss-like Ferns ; the leaves, with the exception of the veins, are most frequently 
formed of only one layer of cells, and consequently stomata are wanting ; the 
formation of the prothallium also somewhat resembles the Mosses. Sori 
marginal, on the extremities of tlic vascular bundles, and surrounded by a 
cupular indusiitni. The sporangia are sessile, with equatorial annulus. Hij- 
meftophyllum (H. (unlridi/ense, European). Trichomanes ('/'. speciostan, Euro- 
pean). Species about 200, which live especially on rocks and trees in damp 
and shady tropical forests. Some have no roots. 

Order 3. Cyatheaceae. Annulus cuntplete and oblique. To this order be- 
long, principally, the tree-like Ferns witli palm-like habit. The number of 
species is about 200, they are all tropical and form forests in some regions of 
Australia. Cibotium and Diclcsonia have marginal sori, with cupular, basal 
indusium. (The stem of D. anlarctica is covered with aerial roots.) Alsophila 
(without indusium) ; djatlica with cupular, inferior indusium (Fig. 212 E). 

FIG. 213. Gleichenia : A part of a leaf with sori ; B a single sorus. 

Order 4. Gleicheniaceae. Sporangia with equatorial auuulus, and longi- 
tudinal dehisceuce, most frequently groups of 3-4 in sori without indusium (Fig. 
213). Gleichenia: the apical growth of tlie leaves continues for a long time. 

Order 5. Schizaeaceae. Annulus apical. To this order belongs Aiu-iinta, 
which is so commonly cultivated in conservatories. The two lowest pinme are 
metamorphosed, having no leaf parenchyma and being covered with sporangia. 
Schiziea. Mohria. Lygodium, a climber, whose leaves have unlimited growth 
and attain a length of several metres. About 70 species. Tropical. 

Order 0. Osmundaceae. The sporangia have at the apex a lateral group 
of strongly thickened cells, which gradually pass over into the ordinary cells. 
The sporangia open by a longitudinal cleft. Indusium wanting. Osmanda 
bears the sporangia upon peculiar, branched pinmu, without parenchyma (the 
uppermost in the leaf). O. reyalis (Royal-Fern) : European. 

Sub-Class 2. Hydropterideae (formerly Rhizocarpre), 

Water Ferns. 

The following further characteristics must be added to those 
given on page 205 : 



Sexual generation. The MICROSTORES produce an extremely 
rudimentary prothallinm, formed of only a single cell, and having 

A B C D 

FIG. 214. Sal"inia nntntis : A microsporan^ium with germinating mierospores and pro- 
truding prothaUia (s) ; B a prothallium with thebicellular antheridium (s) growing out of 
the Tnicrosporangium ; C the two cells of the antheridium have opened by transvetse 
clefts ; beneath is seen the mierospores enclosed by the hardened mucilage ; D spermato- 
zoids still enclosed in the mother-cells. 

tiiso a very much reduced bicellnlar antheridium with a small 
number of spermatozoid mother-cells in each cell (in Salvinia 4, in 
Marsilia and Pilnlaria 16). In Salvinia the mierospores remain 
embedded in a hard mucilaginous mass (at first frothy) which fills 
up the cavity of the sporangium. The prothallium must therefore 



-So/em ri ,,,il,,s. A, B Female prcthallia,/-/, protruding frcm the maerosprre 
11 end scd in the maprosporangii.m; a; archegonin. C An embryo (x 10) 
H connection with tl.r spm-e (s) : the scutiform leaf; ?>- the subsequent foliage- 
B, ..I which V and o stand singly, ,7-e-r in a whnrl ; r the submerged-leaf ; /-/wing- 
like lobes of the prothallium ; m the foot. 


grow out through this slime and also through the wall of the spor- 
angium (Fig. 214), and it thus terminates in a relatively long 

In Marxilia the microspores are set free from the microsporan- 
gium, and the prothallia, with the antheridia, remain in them 
until the spermatozoids are liberated. The latter are spirally- 
twisted threads. 

The MACBOSPORES, on germination, give rise to a very reduced pro- 
thallium, which in Salvinia bears 3 archegonia ; but, if these are 
not fertilised, the prothallium may continue to grow and become a 
fairly large, green body with several archegonia (Fig. 215 A,B). 
In Marsilia the prothallium is still more reduced, it is enclosed in 
the macrospore, and only bears one archegonium. The archegonia 

FiG. 216. Snlvinia natanf. A An archegcmium, unripe, seen in longitudinal section: 
Ji the neck-cells; k the neck -canal-cells ; c the central cell. I? An open archegonium of 
which the neck-cells have separated off. C An open, old archegonium seen from the top. 

are similar in structure to those of the Ferns, but are smaller, and 
sunk more deeply in the tissue of the prothallium. 

The asexual generation is developed from the fertilised 
egg-cell. It is a dorsiventral, horizontal shoot. In Sahinia it bears 
at first a shield-like leaf, the scutiform leaf (Fig. 215 (7, a), which 
is succeeded by the ordinary foliage-leaves. The young plants 
of Marsilia, likewise, have less perfect leaves in the very early 

The formation of the sporangium is the same as in the Lepto- 
sporangiate Ferns. (The 16 spore-mother-cells originate from one 
central, tetrahedric archesporium.) 

The Hydropterideae are divided into 2 orders, the chief differ- 
ences between them being found in the asexual generation. 



Order 1. Salviniaceae. This order more nearly approaches 
the true Ferns, especially so on account of the form of the indusium. 

A B 

FIG. 217. Sah-inia natans (natural size): A seen from above, floating on the water; 
/>' a portion seen from the side in its natural position in the water. 

Only one species is found in Europe, Salvinia natans (Fig. 217). 
Tins is a small, floating, annual, aquatic plant, entirely destitute of 
roots. The dorsiventral, horizontal stem bears two kinds of leaves, 
which are arranged in whorls of three. " Two of these which turn 
upwards are oval, entire, " aerial foliage-leaves " (Fig. 21 7 7>, b 2 -b 3 ) ; 

the third, the "water-leaf" (t* 1 ) is 
submerged and divided into a 
number of hair-like segments, 
similar to the submerged leaves 
in many aquatic plants, for in- 
stance, Water-buttercup (see also 
Fig. 215 C). The whorls of leaves 
alternate with each other; there 
are thus 4 rows of dorsally-placed 
aerial leaves, and two rows of 
ventrally-placed submerged leaves. 
The sporangia are situated in sori, 
each sorus being borne on a small 
column (receptacle or placenta) and enveloped by a cupular, 
but <////'/Wy clnv.d indusium (Fig. 218). The sori are situated on 

I n.. -1 1\ Sori of Salriii/a in longi- 
tudinal section : h. microsporangia; /i 

ni:irti.spMr;iiiLri:i. ( ,-. in.) 



the submerged leaves (Fig. 217 7?, ss~) and are unisexual, i.e. each 
sorus contains microsporangia only, or macrosporangia. 

Azolla belongs to this order. It is a very small, floating, tropical water-plant 
(America and East India), with horizontal, root-bearing stem. The stem brandies 
profusely by lateral buds, and bears the two rows of leaves on its dorsal side, 
the roots on the ventral side. Each leaf is bifi.l, aud divided iutoan upper dorsal, 
ami a lower ventral portiou. The upper segments float on the surface of the 
water and are arranged like tiles on a roof, each one overlapping its neighbour. 
In each floating segment a large cavity is found, in which Annlccna is always 
present. The lower segments are submerged. 

Order 2. Marsiliaceae. The characteristic feature of this 
order, and one not possessed by other Fern-like plants, is that, the 
sori (2-many) are enveloped in leaf- 
segments which close round them and 
form a " sporocarp," just in the same 
manner as the carpels, in the Angio- 
spermous Flowering-plants, close round 
the ovules and form ovaries. The sori 
contain both micro- and macro-sporangia. 
When the spores are ripe, the sporocarp 
opens in order to disperse the spores 
(Fig. 220). 

The two genera (with 57 species, 
Temperate, Tropics) are land- and marsh- 
plants, whose dorsiventral, creeping stem 
bears roots on the under surface, and the 
leaves in two rows on the upper side 
(Figs. 219, 221). The leaves of Mar- 
silia are compound, and divided into four 
small leaflets springing from the apex 
of the petiole (Fig. 219), and resemble 
the leaves of Oxalis. In the bud the 
leaves are circulate (Fig. 219 ?;), and at 
night they exhibit the well-known sleep- 
movements. The sporocarps are borne 
on the petioles of the fertile leaves, near 
their bases (Fig. 219 /) ; they are oblong 
and resemble small beans, the outer cells 
being hard and sclerenchymatous, while FIG. 219. Mantua 

the inner ones are divided into a number (^curalsize): K terminal bmi ; 

o leaves ; / sporocarps ; ,<; point 
of loculi arranged in two rows. On of branching of petiole. 



germination, water is absorbed, the two sides separate slightly, as 
valves (Fig. 220 A), and a long vermiform mass of gelatinous, 
parenchymatoua cells (Fig. 220), swollen by the water, emerges, 
bearing a large number of sori arranged pinnately. Each sorus 
(.<??) is covered by a thin indusium. (The thin covering may be 
considered an indusium physiologically, though not morpho- 

Manilla qiiadrifolia, in Europe. Many species are found in Australia. The 


FIG. 220. Morsi'Hn salaatri.t : A the 
f-porocarp commencing- to germinate ; B 
a more advanced stage of germination. 

FIG. 221. Pilitlarin globulifera (natural 
size) : s sporocarps ; b leaves ; fc tbe 
growing point; r roots. 

ps 01 j/. saivatrix were the means of saving the Burke ex- 
:lor of Australia, and hence this species has earned its specific 

nutritious sporocarps of M. salvatrix were 

pedition in the interior 


'Pilularia has linear leaves, without lamina. The sporocarps are 
spheroid (Fig. 221), brown and hard, and situated near the base of 
tlu- leaves. They are 2-4 chambered and open by a corresponding 
number of valves. 



Class 2. Equisetinse (Horsetails.) 

The characteristics of this class have been described on pao-e 

It is divided into two sub-classes : - 

1. THE ISOSPOROUS EQDISETIN^;. To this sub-class belong, with 
certainty, only the EQUISETACE^E now existent, which are repre- 
sented by only one genus, Equisetum. 

2. The HETEROSPOROUS EQUISETIN.E. Forms which are now ex- 

Sub-Class 1. Isosporous Equisetinae. 

Order. Equisetaceae (Horsetails). 

The sexual generation. The prothallinm is green and leaf- 
like, as in the majority of Ferns, but irregularly branched and 

FIG. 222. Equisetum arvense. The prothallium highly 
magnified. A Male ; s, s aiitberidia. B Portion of a 
female, cut through vertically ; as ae archegoir'a, the central 
one is fertilised ; h h root-hairs. 

FIG. 223. JErQuisetum max- 
imum. Spermatosoids : a 
shows them still enveloped 
by the mother-cell. 

curled. It is often unisexual. The male prothallia bear antheridia 
only, and are smaller and less branched (Fig. 222 A) than the 
female; the latter may attain a diameter of | an inch, and bear 
archegonia only (Fig. 222 B). The antheridia and the archegonia 
resemble those of the Ferns, but the spermatozoids (Fig. 223) are 



larger and less twisted. On the last curve is situated a more or 
less elongated appendage of cytoplasm (Fig. 223 r). 

The asexual generation. The embryo is similar to that of 
the Ferns. The fully developed Eqnisetum is a perennial herb, 
with widely creeping (in some species tuberous) rhizome, from 
which extend erect, aerial, most frequently annual shoots. 

The vegetative aerial STEMS are divided into a number of inter- 
nodes by the whorls of leaves (Fig. 224). The internodes are hol- 
low, the cavities being separated from each other by the transverse 

FIG. 221. Equitetum arvem/e : a fertile branch vrith cone; b vegetative shoot ; c cone ; 

d sporophylls. 

partitions of the solid nodes. The lower portion of the internode, 
which is encased by the leaves, has much thinner and softer cell- 
walls, so that the stem is easily separated into segments just above 
the nodes. Each internode has a large number of ridges and 
tin-rows, and bears at its apex a whorl of leaves whose number and 
position correspond to the ridges of the internode. As in the 
case of other verticillate plants, the whorls are placed alternately, 
one above the other ; the same arrangement is also found in the 
ridges on two successive internodes. In addition to the large 
air-cavity in the centre of each internode (the central cavity), a 


whorl of tabular air-passages is found in the cortex of the stems, 
opposite the furrows (vallecular canals). There is also a similar 
air-passage (carinal canals) in each of the vascular bundles, which 
are placed in a ring, one opposite each ridge, and therefore alter- 
nating with the vallecular canals. The vascular bundles are col- 
lateral as in the majority of Flowering-plants, but poorly deve- 
loped. The xylem of each bundle consists of two groups of 
annular or spiral vessels, close to the outer border of the carinal 
canal, and two groups of scalariform tracheids, each placed on a 
radius passing through a group of spiral vessels. The phloem is 
placed between these four groups, each of which has only a few 
vessels. The stiffness of the stems is mainly due to the large 
amount of silica in the cell-walls of the epidermis, and to the 
sclerenchymatous cells of the ridges. 

All LEAVES are situated in whorls. The VEGETATIVE are simple, 
undivided, 1-nerved, and are united into toothed sheaths (Fig. 
224 a, 1). The branching of the stems in some species (E. arvense) 
is very abundant. The branches break through the base of the 
leaf-sheaths (Fig. 224 b), and generally alternate with the teeth 

The FERTILE LEAVES (sporopJu/lls) are different from the barren 
ones. They are free, shield-like, each one having a short stalk 
bearing usually an hexagonal plate (Fig. 224 d), and closely com- 
pressed into an ear or cone (Fig. 224 a, c). The Equinetums thus 
present an advance in development distinctly beyond that of the 
Ferns, which is further emphasized by the circumstance that a 
transition from the sheath- leaves to the fertile-leaves is found in 
the involucre or annulus, a " collar " of specially modified leaves 
situated at the base of the cone (Fig. 224 a and c). The cone 
may be considered as a very rudimentary flower, and the annulus 
may be regarded as a very early stage in the formation of a, flower 
(perianth). See page 235. 

The SPORANGIA are situated on the underside of the sporophylls, 
one at each angle ; they are sac- like, and open inwardly by a 
longitudinal cleft (Fig. 224 d). An annulus is wanting; but in 
the wall of the sporangium, as in the pollen-sacs of the Flower- 
ing-plants, a layer of cells, with annular or spiral thickenings, is 
developed, which assists in the dehiscence of the sporangium. 

The SPORES are green; the walls composed of four distinct 
layers, of which the outer is gradually separated, except at one 
point, and becomes split into four long bands (elaters) (Fig. 225). 


The elaters are extremely hygroscopic, coiling round the spore 
when moistened, and expanding as soon as dry, presenting a most 
lively object under the microscope when breathed upon and allowed 
to drv. The second layer, when germination commences, becomes 
detached from the inner wall, which is formed of the exospore 
and endospore. 

The order has become much reduced, and at the present time 
includes only one genus, Equi'setnm, with about twenty-five species, 
which are distributed over the entire globe, particularly in damp 
situations. In SOME SPECIES the barren shoots are green and very 
much branched, but the fertile ones are unbranched, pale brown, 
and possess no chlorophyll (E. arvense, Field-Horsetail, Fig. 224, 
and E. maximum}. IN OTHERS the fertile and barren shoots are 
alike green, and either both unbranched (E. luemale'), or branched 
(E. palustre, E. limosum, etc). The fertile shoots of E, silvaticum, 


FIG. 225. -Spores of Equisctum : A damp, with elaters (c) coiled round the spore ; B dry, 

with elaters expanded. 

up to maturity, resemble those without chylorophyll of E. arvense, 
but after that period they produce green branches, and thus re- 
semble the barren ones. 

EXTINCT ISOSPOROUS EQUISETIN.E. In addition to several true species of 
fossilized Equisetums, the order of the CALAMITES, which no doubt is closely 
allied to the Equisetinae, is also found in the fossil state. These were gigantic 
forms, attaining about twenty times the size of those of the present day, and 
stems of nearly 10-12 metres in height are known. They reached the culmina- 
ting point of their development in the Carboniferous peiiod, and died out 
towards the close of the Palaeozoic. The stems had hollow internodes and 
a ternating grooves, similar to their relatives of the present day. The leaves 
must either have been absent or very perishable, since they have not been 
identified with certainty. If the determinations of certain remains of cones 
which of late have been discovered are correct, they were heterosporous and 
had two kinds of sporangia as in Ihe following sub-class. A cambium forma- 
tion and an increase in thickness has been found in the stems. 

Their VSES are very limited. A few species, such as E. hirmale are used for 
polishing on account of the hard siliceous cell-walls of the epidermis, found in 
all species of Etjuisetum. 



Sub-Class 2. Heterosporous Equisetinae. 

The two orders which come under this head are united by the charcteristics, 
that the verticillate leaves are not united into sheaths (Fig. 226), and that be- 
tween each whorl of fertile leaves there is also a whorl of barren ones. The 

FIG. 226. A. fragment of Annularia. 

fertile whorls in ANNULARIES are situated about midway between the barren ones 
(Fig. 227), but in ASTEROPHYLLITE.E they occur immediately above a barren 
whorl (Fig. 228) and contain only half as many members as the latter. The 
lower whorls bear macrosporangia with one macrospore, the upper, micro- 
sporangia with many microspores. 

FIG. 227. Fragment of Annularia longi- 
folia, with sporangia ; the leaves have 
partly fallen off : a barren whorls ; s fer- 
tile whorls. 

FIG. 228. Fragment of cone of Astcro- 
pltyllites (Volltmannia elongata) : a and s 
as in Fig. 227. 

The ANNULABI.E were distichous (Fig. 226), and presumably floating plants. 
The ASTEKOPHYLLITE.E had verticellate branches. These also died out after the 
Carboniferous period, at the close of the Paheozoio. 

W. B. 


Class 3. Lycopodinae (Club-Mosses). 

The characteristics of this class have been given on page 205. 
It consists of two sub-classes, one embracing isosporous, the other 
heterosporous forms. 

Sub-Class 1. Lycopodieae (ISOSPOEOUS Lycopodinae) . 

One kind of spore. Prothallium large, partly green. Leaves 
without ligule. 

Kin. 22'J. Lycopodium annotinwn : A em- FIG. 230. Lycopodiitm clavatum : 

b.vro (n:it. si/.c), with prothallium (pr), one portion of astern, bearing cones (a) ; 

embryo is broken off; B the prothallium ., a spore ; h sporangium in the axil 

(slightly iiiii-.'nified) ; C section through the of a leaf, s. 
i rothallram :nnl embryo in the direction a-b 
of A, iiinl vertically in the plane of the paper. 

Order 1. Lycopodiaceae. The VKOTHALLIUM is only known in 
:i IV\v spi'cics at present, but in these it is more or less tubercular, 
iiiid hears both antlicridia and archegonia. 

In L. iiiii/i'tiinnii the protlialliuin is a relatively large mass of cells, without 
chlorophyll, ;m<l subterranean, in which the antheridia and archegonia are 
embedded 1 ig. :>_".)). In the widely distributed tropical species, L. cernuum, 



and in L. inundatmn, it is a small tubercular body which has a subterranean 
portion, with either little or no chlorophyll ; and an aerial green portion. 
The prothallia of L. phlegmaria and others live saprophytically in the crevices 
of the bark of trees ; they are partly filamentous, branched, and possess no 

The asexual generation. PERENNIAL PLANTS. The stem 
branches monopodially (often apparently dichotomonsly), and is 
thickly covered by small, simple, triangular or scale-like leaves. 
The leaves are spirally arranged in some species (Figs. 229, 230), 
and in others, whose stem is compressed with unequal sides, oppo- 
site (Fig. 231). The roots of Lycopodium are dichotomously 

The SPORANGIA in I/ycopodium are situated singly at the base of 
the leaves, almost in their axils ; they are reniform, unilocular and 

FIG. 231. Lycopodium complanatum : a 
leaves on the edges of the stem ; d leaves on 
the sides. 

FIG. 232. Lycopodium clnvntum. A 
tetrahedral spore seen from above, 
where the three borders join ; and a 
tetrad of bilateral spores, still lying 
in the mother-cell. 

open like a mussel-shell by two valves (Fig. 230 h~). The sporangia 
are developed from a group of surface cells. The archesporium is 
formed from one hypodermal cell (or perhaps a cell-row). 

The fertile leaves are collected upon definite regions of the stem. 
They are either similar to the barren ones, and then the fertile 
portions of the stem pass gradually, without any break, into the 
barren portion (L. selago) ; or they differ from the barren leaves, 
and are then collected into special apical cones (Fig. 230 a). The 
SPORES are tetrahedral or bilateral (Fig. 232). 

About 100 species, chiefly tropical. 

Five species of Lycopodium are found in Great Britain. L. claratum and L. 
selago are common in mountainous districts. L. annotinum is common in the 


Highlands of Scotland. The other genus of the order is Phylloylosstini, with 
one species, P. drnmmondi (Australia, Tasmania, and New Zealand), a small 
plant only a few centimetres high, with two tubers, and about eleven linear 
leaves at the base of the stem which is terminated by a cone of sporophylls. 
FOSSIL Lycopodiacefe in the Carboniferous period. 

OFFICINAL: "Lycopodium," the spores of L. dnvatuni. 

Family 2. Psilotaceae. The sporangia are placed on the apex of short, two- 
leaved stems, as 2-3, seldom four, small capsules. Small herbs, with angular 
stems ; leaves small, simple, and one nerved. Only four species. Psilotnm 
(Madagascar, Moluccas, Sandwich Islands, etc.) is destitute of roots, their place 
being supplied by special underground stems which bear a few modified leaves, 
very much reduced, especially when buried deeply in the soil. Three species. 
Tmesipteris (Australia), one species. 

Sub-Class 2. Selaginelleae (HETEROSPOROUS Lycopodinse). 

Micro- and macro-spores. The prothallia are very much reduced, 
especially the male ; the female does not leave the spore. The 
leaves are ligulate. 




Kn;. L'315. Germination of the microspores of Sclaginclla : A the spore rendered 
transparent, seen from above. In the interior is seen the protballium (/), and the first 
divisions of the antheridium (<i, f>, c, d) ; in B the spore-wall is removed and all 
the spermatozoid-mother-cells formed; in C, the microspore has opened and the spermato- 
zoids and the mother-cells are escaping together. 

The sexual generation. In the MICROSPORES are formed: 
(1) a very small "vegetative" cell, representing the vegetative 
part of the prothallium (/ in Fig. 233 A, B), and (2) a cell many 
times larger and which divides into a number (4-8) of primordial 
cells, each of which divides into four spermatozoid-mother-cells, 
1 hough all of these may not develops spermatozoids. On germina- 
i ion, when the spore- wall is ruptured, the spermatozoids and 
spermatozoid-mother-cells are ejected into the water. 

The SPERMATOZOIDS in Selagmella are elongated and club-shaped, 
with two cilia (Fig. 234) ; but in Isoetes lacustrls they are spirally- 
twisted threads which differ from all other spermatozoids by 



having a bunch of cilia at each end; the other species of Isoiites 
have cilia only at the anterior end. 

The MACROSPORES. Shortly after 
the macrospoi-es have been set free, 
or in Selaginella, while still en- 
closed in the sporangium of the 
mother-plant, they germinate and 
soon become filled with the cellular 
tissue of the prothallium, and even 
in Selaginella the archegonium be- 
gins to be formed before the rupture of the spore-cell-wall has 
commenced (Fig. 235 A'). 

The ARCHEGONIA are constructed on the same plan as those of the 

FIG. 234. Spermatozoidsof Selaginelln: 
b with a remnant of cytoplasm. 


FIG. 233. Macrospore of Selaginella: A longitudinal section, before the rupture of the 
wall, six weeks after being sown. The endosperm (e) has not yet filled the entire 
chamber. Cell-formation is still proceeding in the lower part of the spore. The endosperm 
and prothallium (//)are separated by a distinct line (diaphragm). B Germinating macro- 
spore seen from outside : s wall of the spore ; ce archegonia. 

FIG. 238. Archegonia of Selaginella: A unripe, in longitudinal section ; c the central 
cell; fr neck-canal-cell, which is wedged in between the two-storied neck-cells; B ripe; 
u ventral canal-cell ; C seen from above, open. It will be noticed that the neck is formed 
of two tiers of four cells each. 



other Archegoniatre, but are quite embedded in the prothallium 
(Figs. 235 cr, 236). 

The asexual generation varies very much in the different 

Order!. Isoetaceae (Quill- worts). The only known genus, 
Isoetes (Quill-wort), has an extremely short, tuberous, unbranclted 
stem with very short internodes (Fig. 237). The STEM is 
vemai'kable as being- the only one among the Vascular Cryptogams 
which increases in thickness (see page 202). The meristematic 

FIG. 237. Isoetts lacustris (slightly diminished): st the stem ; r roots ; b leaves. 

FIG. 238. Isoetes lacustris. Longitudinal section through the base of the leaf with a 
microsporangium. The edge of the groove, in which the microspangium is placed, is 
continued as a thin covering which envelopes the sporangium. The inferior edge of the 
ligular groove (L) forms a lip ( J) ; f, sterile cell-rows (trabeculte) which divide the 
sporangium into compartments; I vascular bundle. 

FIG. 239. Selaginella inaqualifolia. Cone in longitudinal section; microsporangia are 
seen on the left side, macrosporangia on the right (most frequently each with four 

cells are situated round the axial cylinder, and form, especially, 
parenchymatous tissue in two or three directions, giving rise to 
- M grooves in which the dichotomously-branched ROOTS are 
produced. The LKAVKS are arranged spirally in a close rosette. 


They are awl-shaped and have at the base a semi-amplexicaul 
sheath, with a groove (fovea), in which a sporangium is situated 
(Fig. 238). The ligule is a foliar outgrowth from the upper edge 
of the groove. The MACROSPORANGIA (each with a number of 
macrospores), are situated on the outer leaves, the MICROS PO RAM ;i A 
(Fig. 238), on the inner ones. Between each cycle of fertile 
leaves there are a number of imperfect or barren ones as in the 
case of the female plant of Cycas. The spores are liberated by the 
decay of the sporangium. The two kinds of sporangia develope at 
the commencement in the same way. The archesporium is, at 
first, a hypodermal layer of cells which grow out in the direction 
perpendicular to the surface of the leaf, and divide by a number of 
walls parallel to this direction, forming a sporogenous mass of 
cells. Some of the cell-rows of this sporogenous mass lose their 
rich protoplasmic contents, and are arrested in their growth; thus 
incomplete divisional walls of sterile cells, " trabeculw," arise in the 
sporangium, dividing it into a number of compartments one above 
the other (Fig. 238 t). (The trabeculae, according to Groebel, play 
the same part as the nutritive cells of the sporangium of Riella 
the tapetal cells, as in the Ferns, are in a great measure dissolved 
at a later period.) The sporogenous cell-rows, in the micro- 
sporangia, give rise to a large number of spore-mother-cells, bufc in 
the macrosporangia only one spore-mother-cell, with tapetum, is 
developed from each fertile archesporial cell. 

The two native species, and several others, are aquatic plants, 
the remaining species are land plants, or are amphibioiis. About 
50 species. In temperate and tropical regions. -FotssiL species 
in the Tertiary period. 

Order 2. Selaginellacese. This order contains only one genus, 
Selaginella. The STEM, in the majority of species, is dorsiventral, 
long and slender, and apparently branches dichotomously, but in 
reality moiiopodially, with well developed lateral shoots. The 
LEAVES are small, round, or ovate, in the majority of species 
arranged in whorls of two leaves each ; these whorls, however, 
are not decussate, but are considerably inclined towards each 
other, an arrangement by which four rows of leaves are produced, 
each whorl having one large and one small leaf. The two leaves 
in each whorl are of unequal size, the smaller one being placed 
on the upper surface and the larger on the lower surface of the 
stem (Fig. 240). Some species have spirally-arranged leaves, more 
resembling the arrangement in the Lycopodiums. 



The FERTILE LEAVES most frequently differ from the barren ones, 
and are collected into spike-like cones (a kind of flower; Fig. 289). 
Micro- and macro-sporangia are found in the same cone (Fig. 239). 
Kiioh sporangium arises from a group of superficial cells of the 
stem, directly over the leaf on which it will be situated later on. 
Bach sporangium has a hypodermal, unicellular archesporium, 
ju.d contains a layer of tapetal cells; these are dissolved later, 
when the spores are ripe, and not before as in the Ferns. In the 

very early stages of their development, 
the micro- and macro-sporangia are 
precisely similar, and the differences 
between them arise later on. In the 
microsporangium all the spore-mother- 
cells divide, and each forms four tetra- 
hedrically-arranged microspores (Fig. 
204) ; but in the macrosporangium 
only four macrospores are formed, by 
the division of a single viotlter-cell, 
while the remaining spore-mother-cells 
are aborted. It is rarely that the 
macrosporangia contain 2 or 8 macro- 

For the GERMINATION OF THE SPOBES, see pages 228, 229. The prothallium 
arises in the macrospore (/-/, in Fig. 235.4), probably by division of the meniscus- 
shaped protoplasmic mass, which is marked off at the apex of the spore ; 
primordial cells are thus formed which later on are surrounded by a cell-wall. 
In six to seven weeks after sowing, the spore-wall is ruptured by the growing 
prothallium, which already has developed archegonia (Fig. 235 a-te). The 
prothallium so formed does not occupy the entire cavity of the spore, but four 
to five weeks after sowing, the large-celled parenchyma is developed in the 
lower portion of the spore by free cell -formation; this has been termed by Pfeffer, 
" endosperm." since it is similar to the endosperm of Flowering-plants. Goebel, 
however, has termed it " secondary prothallium," as the homology with the 
endosperm of the Angiosperms is very doubtful. 

The IT. UTILISED opSPHERE divides into an upper (hypobasal) and 
;i lower (epibasal) cell; from the latter alone the embryo is deve- 
loped with its root, stem, foot, and two cotyledons, and the former 
gives rise to an organ which appears in this instance for the first 
time, but which occurs in all Flowering-plants, viz. the suspensor. 
This forces the embryo down into the "endosperm," which is 
entirely or partially absorbed by the embryo. In the case of the 
Flowering-plants the embryo is developed with its longitudinal 

Fit;. 240. Selaginella martens!! 
.- lower leaves; r upper leaves. 


axis in the elongation of the suspensor, but in Selaginella the em- 
bryo is situated transversely to it. 

SelagineUa (300-400 species), is essentially tropical, only one 
species living in the North (S. spinulona) , but others grow in 
Central and South Europe. 

Order 3. Lepidodendraceae are extinct, tree-like Lycopods, which are 
found especially in the Lower and Middle Carboniferous. Vegetatively they are 
most nearly related to Lycopodium, but the stem attained much larger propor- 
tions (about eleven metres in height and one metre in thickness), and had a 
cambium by which it increased in thickness. It was regularly dichotomous, 
and closely studded with spirally-placed leaves, which left behind them peculiar 
rhombic scars. The large cones resemble Pine-cones, and bore sporangia 
much larger than any which are now produced (the male ones as much as 
2 cm.'s in length). The macrosporangia were situated at the base, and the 
microsporangia at the apex. 

Order 4. Sigillariaceae. These are, presumably, another group of extinct 
tree-like Lycopods (especially in the Middle Carboniferous). The name has been 
derived from the seal-like scars, which the fallen leaves have left behind in 
longitudinal rows on the grooved stem. The rhizomes of these plants were 
formerly termed Stitjinariu, and placed in a separate genus. 

Order 5. Sphenophyllaceae form an entirely extinct group. They do not 
definitely belong to any of the three large classes of Vascular Cryptogams, but 
it is perhaps best to place them in juxtaposition to these. They were herbaceous 
plants with verticillate, wedge-shaped leaves, with nerves branching dicho- 
tomously into equally strong branches. Micro- and macro-sporangia were 
formed in the same cone ; and were situated in the axils of the leaves, as in the 

The Transition from the Cryptogams to the 

All the plants considered in the preceding chapters are included 
in the term CRYPTOGAMS; all in the following chapters under the 
head of PHANEROGAMS (see page 3). Hoffmeister's pioneer works 
(Ibol, Vergleichende Untersuchungen der hiJhereu Kryptogamen, etc.) 
and the numerous researches published later by other investigators, 
have closed the gap which was formerly thought to exist between 
these plants ; so that we now, in the series : Bryophyta Pterido- 
phyta Grymnospermse see the expression of a 
single line of development in accordance with a definite plan. The 
forms through which this gradual development has taken place 
have in course of time, however, to a great extent died out, and 
only single links of the chain connecting the lowest to the highest 
still remain. 

THE ALTERNATION OF GENERATIONS, which we found indicated in 
certain Thallophytes, can be proved with the greatest clearness in 
all the higher Cryptogams, from the Mosses upwards; it is also 
found in the Phanerogams, but not in such a pronounced degree, 
because one of the generations is so far reduced that it has almost 
given up its independence. For the sake of greater clearness, we 
will begin with the comparison of the sporophyte, asexual (second) 

The asexual (and) generation of the Cormophytes. 

The asexual generation which follows from the further develop- 
ment of the fertilised oosphere, is, in the Mosses, only the sporo- 
gonium (according to one theory it is perhaps homologous with a 
spore-bearing leaf, situated upon a short stem, see p. 187) ; in 
Filicince, Equisetince, and Lycopodinw, on the other hand, it is a 
highly developed plant differentiated into stem, leaf, and true root, 
and bearing the sporangia on its leaves. The MODIFICATION OF THE 
s IK mi is very slight in Filicmw. The first leaves of the embryo 
;m- very simple in form (Fig. 205), but after a certain age all the 
leaves which arise are essentially alike. The fertile leaves do not 



differ from the barren ones, and are found associated with them, 
and their formation does not limit the growth in length of the 
stem. It is only in a few of the true Ferns, and in the Hydro- 
pteridese, that the fertile leaves differ considerably from the 
barren ones. A division of labour in which certain leaves are set 
apart for nutrition, and others for reproduction, is found more 
pronouncedly in the Eqnisetime and Lycopodinw, for in these 
groups, with a few exceptions, the fertile and barren leaves are 
very dissimilar ; the former are collected in special ear-like cones, 
which terminate the further growth of the short stems on which 
they are borne. In connection with the cone, leaves are sometimes 
developed which form a transition from the barren to the fertile 
ones (the " annulus " in Equisetaceoe), and in these cases the first 
indication of a flower with perianth or floral-leaves is to be traced. 
Among the Cryptogams the highest division of labour is found 
in Selaginella and Isoetes, which have the two kinds of sporangia 
borne on different leaves. The division of labour (modification) 
is, however, still more pronounced in the Phanerogams : the leaves 
which bear the micros porangia (" pollen-sacs ") have quite differ- 
ent forms from those which bear macrosporangia (the " nucellus " 
in the ovule), the former are termed stamens, the latter carpels ; 
in certain instances, too, there is even a contrast between the 
" male plants" and the " female plants." Moreover, a division of 
labour, in a much greater degree, takes place in the leaves which 
do not directly take part in reproduction, and it is thus possible 
in many plants to draw a sharp line not only between stamens and 
carpels, but also between four or five distinct kinds of leaves, which 
differ in form, structure, and corresponding functions, and which 
appear in regular sequence on the shoot: namely, between " scale- 
leaves" and "foliage-leaves," 1 both of which occur in the Crypto- 
gams, and the " floral-leaves," including the bracts and leaves of the 
"perianth," which latter often differ from each other in form and 
colour, and are then separated into sepals and petals. The leaves 
stamens and carpels which bear the sporangia are termed sporo- 
phylls, and the shoot, or extremity of a shoot, whose leaves are 
modified into sporophylls, is terminated in its further growth by their 
production, and is knoivn as a jiower. The flower which is most 

1 Floral-leaves (hypsophyllary leaves) are here adopted as an equivalent of 
the term " Hochbliitter," to signify leaves on the floral-shoot other than foliage 
or sporangia-bearing leaves. The term bract is applied only to leaves in whose 
axil a flower is borne, and bracteoles to leaves borne on the flower-stalk (pedicel}. 



completely furnished has calyx, corolla, stamens, and carpels ar- 
ranged in this order. If the only sporophylls present are stamens, 
then it is said to be a male (staininate) flower, and if only carpels, 
then a female (pistillate) flower, and in both these cases the flowers 
are nni*c,cnal, or diclinous. If stamens and carpels are both present 
in the same flower, it is termed hermaphrodite. Diclinous plants 
in which the female flowers are situated on one plant, and the 
male flowers on another, are termed dioecious ; and those in which 
the same plant bears the two kinds of flowers are termed 'monoe- 
cious. When the male, female, and hermaphrodite flowers are 
found in the same species, the plant is said to be polygamous. 

The sporangia-bearing leaves Sporophylls. In the 
Mosses the asexual generation is only represented by the sporo- 

gonium, and if the theory is 
correct which considers the 
sporogonium to be an em- 
bryo consisting of a rudi- 
mentary stem and terminal 
leaf, then the spores are pro- 
duced on the leaves in these 
plants. The sporangia in 
the Filicinje are situated in 
groups (sori) on the back or 
on the edge of the leaves. 
The number of sporangia in 
the sorus diminishes very 
greatly in the Marattiaceae 
and Gleicheniaceae (three to 
four in the latter, Fig. 213). 
In the Equisetinse the spor- 

l-'iu. 2U. Ci/cns : <i stamen (nat. size) seen 
frnti: tlic uiuk-r side ; b four pollen-sacs, not yen 
open, forming a "eorus " ; c three open pollen, 
sacs ; of a pollen-grain. 

angia are situated in a small 

number on the under-side of 
shield-like leaves, and in 
Lycopodinse, singly, in the 
axils of the fertile leaves, which are alike and bear either micro- 
or macro-spoi-angia. In the Phanerogams there is a great differ- 
ence between the stamens and carpels. 

Stamens. In the lowest Phanerogams (Cycadece) there are 
many inductions of relationship to the Ferns. The stamens are 
flat and broad, and have on the back many pollen-sacs (micro- 
*j>"r<i)i<jin) arranged in small groups (true sori), which even have 



a small " placenta," similar to the one possessed by the Ferns, 
and open towards the inside by a longitudinal cleft (Fig. 241, 
compare Fig. 213). A section of the Coniferce agree more closely 

FIG. 242. Stamens of Araucaria (pollen- 
sacs long and pendulous). 

FIG. 243. Hale flower of Tax us. 

with the Equisetacene, in having a few (three to eight) pollen- 
sacs arranged on the under-side of more or less shield-like leaves 
(Figs. 242, 243, compare with Fig. 224 a, c, d). In the Abietacete 
the number of sporangia is diminished to two, which are placed 
also on the lower side (Fig. 267) of a stamen. The number of 
pollen-sacs (microsporangia) in the Aiigiosperms is nearly always 
four to each stamen ; they are longitudinal projections which are 

C t 

FIG. 211. A Cross section through a qnaclrilocular anther in different stages of develop- 
ment : s the seam where it bursts open ; vf vascular bundle ; fc connective. B A stamen. 
C Another stamen seen from the front (f) and from the back (b). 

placed in pairs on each side of the central line of the stamen, two 
on the edge, and the other two generally on the side which is 
turned inwards ; the pollen-sacs generally dehisce longitudinally 



(quadrilocular anthers, Fig. 244). A few, for instance Orchidaceoe 
and Asclepiadacea?, have only two pollen-sacs (bilocular anthers) ; 
and in others, such as Solanum and the Ericaceae, they open by 
pores ; in Lauracere and Berberidacese, by valves. The part of the 
stamen which bears the pollen-sacs is termed the anther. Most fre- 
quently this is supported by a stalk known as the filament. 

Carpels. The simplest forms of cai'pels are found in Cycas. 
In this genus both the foliage and fertile leaves are pinnate, and 
hence present great similarity ; the ovules (macrosporangia) are 
situated on the margin of the central portion, just as the sporangia 
are placed on the edge of the fertile leaf of Opliioglossum (Fig. 
245, compare with Fig. 209). The carpels of the other Cycadea? 
present greater divergence from the foliage-leaves, being peltate, 
for instance, in Zamia and Cemtozamia (Fig. 246). The ovules 

FIG. 215. A carpel of Cycas rcvoluta with 5 
ovules (s), at half to one-third nat. size. 

FIG. 240. Carpel with 2 ovules of 
Ceratozamia vobiisto. (i). 

in the Coniferse are situated on the upper side and near the base 
of the ovuliferous scales, almost in the same position as the spor- 
angia in the Lyeopodina? (Figs. 269, 272, 273 IT, compare Figs. 
2:111, :j:!'.i). In '!'., * the uninclosed ovule is placed on the apex of 
a shoot (Tig. 261). In all these plants the ovules are not encln^'d 
l>y tin- carpi-Is, that is, they are not enclosed in chambers formed 
by the turning in of the walls of the carpel, and hence the name 


Gymnospermce is given to them. In the higher Flowering-plants, 
the Angiospermce, the ovules are distinctly situated on the edge, the 
upper surface, or base of the carpel ; but the carpel closes round 
the ovules which are therefore enclosed in a chamber the ovary. 
In a few cases, for example in the Polygonacese, an ovule is situated 
apparently on the apex of the stem itself, as in the Yew ; in other 
cases, as in the Primulaceae, many ovules are apparently devel- 
oped on the apex of the stem, which seems to have been speciallv 
adapted as a placenta, but it is also possible and correct in these 
cases to suppose that the ovules are in reality developed on 
the carpels. 1 A single fully-developed carpel or a collection of 
carpels joined together is termed the pistil. The extremity of the 
carpel, which is specially developed to catch the pollen-grains and 
form a suitable nidus on which they may germinate, is called the 
stigma. The united edges of a carpel which bear the ovules are 
termed the ventral suture. The back of the carpel forms the dorsal 
suture. The Marsiliacea? take a position among the Hydropteridere 
analogous to that occupied by the Angiosperms; the sporangia are 
in a corresponding manner enveloped in a closed leaf. 

The collection of stamens in a flower is termed the androecinm, 
and all the carpels, whether individually free or united into one 
pistil, the gynceceum. 

The Sporangia. The asexual generation of the Mosses is the 
sporogonium, in which the spores arise in tetrads from the mother- 
cells. The sporangia in the FiUcince take their origin either from 
a single cell (Leptosporangiata?) or, what probably may be regarded 
as an older stand-point, from a group of cells (Eusporangiata 1 ). In 
both cases there may be distinguished in a mature sporangium 
three tissues, which have different significance (Fig. 204) : (1) an 
external layer, the sporangium-wall, most frequently composed of 
one layer of cells made up of cells of dissimilar structure, so that on 
desiccation the wall is ruptured and the sporangium opens in a 
definite manner; (2) an internal group of cells, consisting of the 
spore-inother-cells, developed from an archesporium, and which by 
division into four gives rise to the spores ; (3) a layer of cells 
lying' between the two already mentioned, which is dissolved before 
maturity. The intermediate cellular layer, which directly sur- 
rounds the spore-forming cells, is in form and contents more worthy 
of note than the others, and is termed the tapetum. The construc- 

1 It may be here remarked that another explanation is possible, based on the 
study of the development (A") . 



tion of the sporangium in the Equisetince and Lycopodince is in the 

main the same. 

In the PHANEROGAMS the Microsporangia are termed Pollen- 
sacs. They take their origin from a large group of cells, which, in 
the Angiosperms, lie immediately beneath the epidermal cells of the 
anther. In the developed, but not yet mature, sporangium (pollen- 
sac) there are to be found: (as in the Vascular Cryptogams) (1) an 
internal group of mother-cells which give rise to the pollen-grains 


FIG. 247. Development of an anther. .4 Transverse section of a young; anther of Doroiii- 
citm iiKU'roji/iylfmii. The formation of the 4 pollen-sacs commences by divisions of the 
hypoderrnal cells (at m, for instance). These cells divide by periclinal walls into external 
cells which only take part in forming the anther-wall ; and internal cells, which corres- 
pond to the Archesporium, and from which the spores are derived. These spore-forming 
cells are drawn with thicker walls in B-E. The commencement of the vascular bundle is 
seen in the centre. B An older stage ; the pollen-sacs already project considerably. It 
is the cells in the hypodermal layer which are active and in which tangential divisions par- 
ticularly occur ; / v vascular bundle. C A corresponding longitudinal section. D Trans- 
verse section through an older anther, the thickness of the wall outride the mother-cells 
nf the pollen-grains is already increased, and it becomes still thicker by the division of the 
hypodermal cells : its most external layer of cells but one, becomes transformed into the 
" fibrous cells." E Transv r erse section of a still older pollen-sac of Menyanthes ; sm are 
the mother-cells of the pollen-grains surrounded by the tapetnm (t), external to the tapetum 
is the anther-wall, which is still far from being fully developed. The sub-epidermal layer 
becomes " fibrous," and the cells lying inside it become dissolved, together with the tapetum. 

(microstores), in this case also formed in tetrads; (2) a group of 
cells surrounding these, of which the internal ones form a tapetal 
similar to that in the Vascular Cryptogams ; the tapetum 



and some of the cells surrounding it in this group, become dissolved 
before maturity ; the more external ones, on the other hand, are 
provided with peculiar thickenings, and form the " fibrous " layer 
by the aid of which the dehiscence of the anther takes place; 
(3) an external layer, the epidermis, enclosing all the other layers 
(Fig. 247). 

In some Conifers (Ciqwessits, Thuja, and several species of Jitniperux) the 
microsporaugia (pollen-sacs), which are situated on the under side of the stamen, 
are covered by a thin structure which seems to be a continuation of the lamina 
and which is supposed to be homologous with the indusium of the Ferns. 

The Ovule in the Phanerogams arises most frequently on a 
projecting portion of the carpel, termed the placenta. The ovules 
(compare the sporangium of the Eusporangiatae and especially the 



FIG. 248. Development of the ovule in the Red Currant, Ribes nibrum, arranged alpha- 
betically in the order of development. A Is the youngest stage, E the oldest, ii Inner 
integument; ie outer integument ; we nucellusj in archespore (mother-cell of the euibryo- 

pollen-sac) take their origin from a group of cells which lies beneath 
the epidermis (Fig. 248 A, JB). First of all a small papilla is 
formed, which is later on provided with a vascular bundle and 
becomes the funicle ; this probably has the same value as the 
projections ("placenta") on which the sori in the Ferns are 
attached. Only one macro sporangium (nucellus ; Fig. 248 we) 
is developed at the apex of the funicle. This arises by a process of 
cell-division exactly corresponding to that by which the pollen- 
w. B. 



sacs are formed (Fig. 248 C-E), with this difference only, that 
while a great many cells may be distinguished in each pollen- 
sac, which forms pollen-grains by tetrad-division, only a few are 
found in the ovule, and all these moreover are suppressed, with one 
siui/le exception which developes into the macrospore (embryo- 
sac) without undergoing a division into tetrads. The wall of the 
embryo-sac, in the Grymnosperms, may be thick and divided into 
two layers and partly cuticularized, as in the spores of the Cryp- 
togams which are to be set free. In the Angiosperms, on the other 
hand, the wall is extremely thin. 

The pollen-sac thus stands in the same relation to the nucellus 
as the micros porangium does to the macrosporangium : in the 
pollen-sacs and microsporangia a number of spores arise by the 
tetrad-division of several mother-cells ; in the nucellus and macro- 
sporangium, a reduction of the cells already formed takes place to 
such an extent that the number of macrospores becomes one 
(Salvinia, Harsilin, Phanerogams) or four (Selaginella), or rarely a 
large number as in Isolates. 

In the Ferns, as stated on page 210, etc., indnsia covering the 
sori very often occur. Horsetails and Club-Mosses have no 
indusium; but in all Phanerogams cupular or sac-like structures 
(integuments') are found which envelop the nucellus. These de- 
velope from the upper end of the funicle (ii and ie, in Fig. 248 ; 
y and i, in Fig. 249) and enclose the nucellus on all sides as a 

sac, leaving only a 
small channel at the 
apex of the nucellus 
the m icropyle 
(Fig. 249) through 
which the pollen- 
tube proceeds to the 

FIG. 219.-Various forms of ovules: .4 an erect ovule embryo - sac. The 

(orlhotropous) ; -B reversed (anatropous); C curved (campt/7o- ovules of the Grym- 

',-. T oi(s) : fc the nueellus (shaded in all the figures) ; s the em- .ho, m lu 

bryo-sac ; ,-h the base of the ovule (chalaza) ; i, and i the y 

external and internal integuments, the dotted line denotes One integument 

bhe place where the scar (hilum) will form when the seed /T7- , 9^1 Oft t OtfQ 

is detached from the funicle. * lgS ' ^^ ^^ ^ J ' 

274) and the same 

i> the case with the majority of the Sympetalae and a few Cho- 
ripftalsv; but the Monocotyledons and most of the Choripetalte 
have two integuments (Fig. 249). 

In shape the integuments resemble very closely the cupular 



indusinm of the Hymenophyllaceae, certain Cyatheacere (Fig 
212 E), and Salvinia (Fig. 218) ; that they are really homologous 
with these is probable, bat is not proven. Some authorities 
regard them as structures found only in the Phanerogams. 

The ovule is thus a " monanyic " (i.e. reduced to 1 sporangium, 
the nucellus) sorns, situated on afunicle, and enclosed by one or two 
cupular indusia, the integuments. Some of the ovules are erect 
(prtJiotropous) , others curved (campylotropous) , the majority reversed 
(anatropous) (Fig. 249). 

[Goebel (1884 and earlier) with Strasburger considered the entire ovule of the 
Phanerogams as homologous with the macro sporangium, the integuments how- 
ever, as new structures in contradistinction to the Ferns : the funicle then corre- 
sponds to the stalk of the sporangium. The integuments of the ovule (according 
to Goebel, 1882) differ from the indusium of the Fern-like plants in being devel- 
oped from the basal portion of the nucellus and are not, as in the Ferns and 
Isoetes, a portion (outgrowth) of the leaf which bears the sporangia (7iT).j 

The nucellus is the only macrosporangium which never opens; 
the macrospore remains endowed in it, and the macrosporangium remains 
attached to the mother-plant. It is therefore essential that the 
method of fertilisation which is employed should be very different 
from that of the Cryptogams. The pollen-grains must be transferred 
to the ovule, and retained either by a drop of mucilage at thr micro- 
pyle (Gymnosperms) or by the stigma on the carpels (Angiosperms). 
Fertilisation by spermatozoids, which are freely motile in water, is 
abandoned in the Phanerogams. 


Many other modifications, unknown in plants of more simple 
structure, take place, for instance, in the shoots which bear the 
fertile leaves; especially in the form of the stem or thalamus (hypo- 
irynous, perigynous, epigynous) ; in the development of the peri- 
anth which stands in intimate connection with the special means 
employed to effect fertilisation; with respect to the different 
grades of union found in the leaves; in the union of the flowers 
into aggregations of a higher order (inflorescences), and at the same 
time the production of " floral-leaves " (page 235). 

The sexual generation. The Fertilisation. 

The sexual generation in the Mosses is relatively well developed, 
because not only the protonema, but all the other vegetative 
parts of the Moss-plant, in addition to the archegonia and anthcr- 
idia, belong to it. In the groups which follow, a gradual but in- 
creasing reduction of the sexual generation takes place, and at the 


same time an indication of sex is found in the prothallia, which 
finds expression in the forms of the spores themselves. In the 
majority of cases among the isosporons Vascular Cryptogams, the 
sexual generation prothallium is a green, leafy expansion which 
can sustain itself by the assimilation of carbonic acid, and by the 
absorption of nutriment from the soil by means of root-hairs. In 
some plants ( Ophwglossacece, Lycopodium annotinum)tihe prothallium 
is a subterranean, pale, tubercular body, but in these instances it 
is relatively large. In the heterosporons Vascular Cryptogams and 
in the Phanerogams, the prothallium is much more reduced, both 
as regards its size, and also with respect to the number and struc- 
ture of the antheridia and archegonia. 

1. The Microspores. The PEOTHALLIUM in all Vascular 
Cryptogams which have unequal spores, consists of a single, vege- 
tative (barren) cell, which plays a very unimportant part in the life 
of the prothallium (Fig. 233 A*). In Stilriiu'n it is somewhat elon- 
gated and tubular, because it must break through the sporangium 
( Fiy. '214) ; but in other cases it is very small and lenticular. In all 
these plants only one antheridium is formed. In Salvinia it con- 
sists of 2 cells whose walls are ruptured in order that the spermato- 
zoids may be liberated (Fig. 214 7>, C). In Marsilia, Isoiites, and 
SelaijiiK'Ud the prothallium does not leave the spore, and consists 
for the most part of primordial spermatozoid-mother-cells without 
ci-lf -tcnU, which on germination are ejected so that the spei-mato- 
zoids are set free. 

In the Phanerogams, the microspores have from olden times 
leen. termed pollen-grains. 

In the GYMNOSi'KRJis the prothallium is reduced to 1, 2 or 3 small 

cells, placed on one side of the mature pollen-grain (at the top 

in Fig. 250 I, 77, and in Fig. 267 N) and which do not play any 

part in the germination of the pollen-grain. The antheridium is 

represented by the remaining portions of the interior of the pollen- 

<_:rain. that is, it consists of a large cell with a nucleus which does 

not even go so far as the antheridium of SelagineUa and become 

divided into spermatozoid-mother-cells without cell-wall, for even 

these cells are not formed. The unicellular antheridium grows, 

on tin- germination of the pollen-grain, into a tubular body known 

as the pollen-tube, formed from the inner wall of the pollen-grain 

I Fig. -J ">"), which works its way down the micropyle to the 

oosphere. The FIT! ilisation takes place by diosmosis through the 

cell-wall, and consists here also of the coalescence of the nucleus 



of the pollen-tube (the sperm-nucleus, male pronuclens) with that 
of the oosphere. 

In the ANGIOSPERMS the reductions proceed still further. The 
barren cell, which represents the prothal- 
lium, was in the last group separated from 
the antheridium by a true cell-wall, but in 
the Angiosperms a membrane at most, but 
no firm cell-wall, is formed. The pollen- 
grain contains two cells, a vegetative and a 
free generative cell. Both these pass into 
the pollen-tube, but the vegetative cell dis- 
appears about the time the pollen-tube 
reaches the ovule ; while the generative 
cell divides into two: one, the sperm-nucleus 
coalescing with the nucleus of the oosphere, 
the other being absorbed (Lilium, after 

The Gymnosperms prove in yet another 
point that they are more closely related to 
the Cryptogams than are the Angiosperms. 
When the pollen -grain begins to germinate 
the external wall ruptures as in the Cryp- 
togams (Fig. 250), but in the Angiosperms special germ-pores are 
formed in the cell-wall for the emergance of the pollen-tube. 

2. The Macrospores. The prothallium in Salviuia and Marsilia 
is still rather large, green, and capable of the independent assimi- 
lation of carbon. It projects more or less from the macrospore and 
bears (in J/iam7ia only one, in tialvinia several) archegonia, which 
however are embedded to a greater degree in the prothallium, and 
are more reduced than the archegonia of the true Ferns and Horse- 
tails (Figs. 215, 216). The prothallium is still more reduced in 
Isoetcs and Seh.ii/iueUa partly because it is smaller and is in a higher 
degree enclosed in the spore, it also contains less chlorophyll, or 
is entirely without chlorophyll, and in consequence incapable of 
independent existence, whilst the number of archegonia is less ; and 
partly because the archegonia are themselves reduced, the cells of 
the neck are fewer and embedded to the level of the surface 
of the prothallium without any, or with only a very slight 
projection (Figs. 235,236). Finally, the prothallium with its arche- 
gonia begins to develope in Selaginella while the macrospore is still 
within its sporangium, and before it is set free from the mother- 

PIG. 250. I Pollen-grains 
of Cupressus ; at the top is 
seen one proth;illium-cell. 
II Germinating ; a pollen- 
tube; a the extine ; li the 



plant. After the spores are set free and germination has com- 
menced, the spore-wall ruptures and the prothallium is exposed. 

The GYMNOsi'ERMSgo still farther. The macrospore (embryo-sac) 
germinates and forms internally a cellular tissue, designated in 
former times by the name of albumen (endosperm), which is homo- 
logous with the prothalUnm. It always remains enclosed in the 
i-mhnjo-sac, and is a parenchymatous mass containing a large supply 
of nourishment. In the upper part of the endosperm a number 
of archegonia are developed which are in the main constructed in 

t - 

FIG. -51. Longitudinal sec- 
tion of ovule of Abies cana- 
iltnsif. Inside the integument 
( ; ) is seen the nucellus, n ; 
in the micTopyle. In the in- 
terior of the nucelliis is seen 
:in oval mass of cells, the en- 
tm, anrl at its top two 
arche^oniii, c. The ovule ia 
turned in such a way that the 
micropjle points upwards, 
but usually it turns down- 
ward in the Abietinfce. 

FIG. 252. The apex of the nucellus () of an ovule of 
Abies: I lonK-^haped cells which guide the pollen-tube; 
s the wall of the macrospore (embryo-sac) ; li the neck- 
cells of the arcbeponinm ; ); the ventral canal-cell; and 
c the central cell (oosphere). The archegonia of the 
Cryptogams should be compared with this (see pages 181, 
208, 216). 

the same manner as those in the Cryptogams, but are still more 
reduced, the neck consisting only of 4, 2, or 1 cell (Figs. 251, 252). 
The ventral canal-cell is also formed, in the majority, as a small 
portion cut off from the large central cell just beneath the neck ; 
the larger remaining portion becomes the oosphere. When the 
pollen-tube has passed down to the oosphei-e (Fig. 253) and fer- 
tilisation has been effected, the oospore commences a cell-formation, 
tin- linal result of which is the formation of an embryo (the asexual 
generation') which is provided with a thinner, lower end, termed 
the suspensor. The embryo is forced mor'e or less into the endo- 



sperm in -which it may rest for a longer or shorter time, and 
generally is developed to such an extent that it has a distinct 
primary-root (radicle) and stem (plumule) with one or more 
embryo-leaves (cotyledons). 

When the oospbere has been fertilised its nucleus sinks down to its lower end, 
and by repeated division into two, forms four cells lying in one plane (Fig. 253, 
see base of the left archegoniurn). Three tiers of cells are now formed by trans- 
verse division of these four. It is the intermediate one of these which elongates 
and forms the suspensor, or four suspensors, if they separate from each otber, 
which push the lowermost four cells deep down into the endosperm. It is 
from these four lower cells that the embryo (or four embryos when the suspen- 


FIG. 254. Embryo-sac of 
Cnrex prcecox : syn synergi- 
d; ; fcb the oosphere; c the 
central nucleus ; ant the 
aaiipodal cells. 

FIG. 253. The apex of the nucellus (n) of Abies in longi- 
tudinal section : c, c the oospheres of the two archegonia ; 
the embryo-formation has commenced at the bottom of 
the left archegonium ; s wall of the macrospore ; p pollen- 
grains ; r pollen-tubes. 

SOTS separate) is developed, but never more than one embryo attains full devel- 
opment. As several archegonia are contained in one and the same ovule, all of 
which are capable' of forming embryos, there is the possibility that several 
embryos may be developed in a seed (po'yembryony), but usually only one 
embryo attains perfect development. 

At the same time that the embryo is being developed, other 
changes are taking- place in the ovule, especially in the integument 
which becomes the shell of the seed (testa'). The endosperm grows, 
and the embryo-sac supplants the cells of the nucellus. The seed 



is now formed, and it consists in its most complete development, 
as in this instance, of three parts : 

(1) The testa of the seed, formed by the enveloping integuments, 
with the remainder of the tissue of the nucellus lying outside the 
embryo-sac (the macrosporangium). 

(2) The endosperm or prothallium. 

(3) The embryo. 

The reduction in the ANGIOSPERMS is carried to the extreme 
limit. In the embryo-sac (the macrospore) the nucleus by con- 
tinued division produces a prothallium consisting of primordial 
cells (Fig. 254). In the upper end of the embryo-sac (which is 
nearest the micropyle) are three cells, two of which are termed 
the "co-operating cells" (synergidcv) and the third is the oosphere. 
Three others are placed at the opposite end of the embryo-sac and 
are therefore termed the ' antipodal cells." Finally, a large cell is 
also formed, which occupies the space between the two groups and 
whose cell-nucleus, the central definitive nucleus, lies in the centre 
of the embryo-sac. These primordial cells are the slight rem- 
nant of the prothallium. The entire structure of the archegonium, 
with its neck and canal-cells, has disappeared, and nothing is left 
but the indispensable oosphere. When the oosphere has been 

fertilised, and has com- 
J I menced the cellular divi- 
sions which lead to the 
formation of the embryo 
(Fig. 255), the synergida> 
and antipodal cells are 
absorbed, and a cell-for- 
mation begins by a new 
process which emanates 
from the definitive nu- 
cleus and by which a pa- 
renchymatous cell-tissue, 
the nutritive-tissue, arises 
which may perhaps be 
considered as homologous 
Avith the endosperm of the 
Gymnosperms. The dif- 
ference is that the nutritive-tissue of the Angiosperms is formed 
in two parts with an intervening interruption ; the primary 
nutritive-tissue is first formed, and after fertilisation is absorbed, 


Fie. 255. Diagrammatic longitudinal section 
through an anatropous ovule shortly after fertili- 
sation ; a. and i are the two integuments ; / the 
funicle ; ): the nncellus; S the embryo-sac, with the 
incipient formation of nutritive-tissue ; E the 
embryo; P the pollen-tube passing through the 
micropyle () to the oosphere. 


with, the exception of one cell, which continues the development 
and gives rise to the nutritive-tissue proper, which is formed in 
the first instance of primordial cells, and later on of a cellular 
tissue ; this nutritive-tissue formed in the embryo-sac is termed 
" endosperm "; in a few instances l a tissue which is derived from 
the nucellus functions as nutritive-tissue, and is termed " peri- 
sperm." In many plants the seeds, when ripe, contain a very 
rich nutritive-tissue, in addition to the embryo, for the purpose of 
its nourishment during germination. These are termed albu- 
minous (endospermous) seeds, in distinction to the ex-albu- 
minous, or those in which the nutritive-tissue is stored in the 
embryo itself, before it is completely developed, and used for its 

In addition to the changes which fertilisation produces in the 
ovule itself, it also gives the impetus to a series of changes in the 
entire shoot which bears the ovule. The perianth, stamens, and 
style, generally wither, because the part they play is at an end ; 
the wall of the ovary grows and becomes the wall of the fruit 
(pericarp). The entire gynoecium of a flower, transformed as a 
consequence of fertilisation, is termed a fruit. It consists of two 
parts, the pericarp and the seeds, and according to the nature of 
the pericarp, the fruit is termed a capsule, nut, berry, or drupe. 

The chief characteristic of the Phanerogams does not lie in the 
formation of the flower (although they may quite properly be 
termed " Flowering-plants "), because Equisetums and Lycopods 
have reproductive shoots as highly differentiated as those of 
certain Gymnosperms and other Phanerogams. As regards the 
SEXUAL GENERATION the characteristics are found : (1) in its 
great reduction ; (2) in the transmission of the microspore (pollen- 
grain) to the macrosporangium, and its germination, with the 
formation of a pollen-tube (antheridium), the protoplasm of which 
is not differentiated into spermatozoids ; (3) in the fact that the 
macrospore (embryo-sac) never leaves its sporangium (nucellus) ; 
and further in the Angiosperms, (4) in the peculiar development 
of the nutritive-tissue in two parts ; and (5) in the great reduc- 
tion of the archegonium. 

As regards the ASEXUAL GENEHATION the characteristic feature is 
that this generation is formed whilst the sporangium is still 
attached to the mother-plant, and for a long time is nourished by 
it ; and that after the sporangium has become detached from the 

1 Piperaceie, Nymplueaceas. 


mother-plant, it spends a longer or shorter resting period as the 
embryo in the seed (enveloped by the testa), and does not make its 
appearance until the "germination " of the seed. In addition the 
shoot -which bears sporangia undergoes greater modification than 
in the case of the Flowerless-plants. 

The Phanerogams are separated into two Divisions as follows : 
Division 4. Gymnospermae. The ovules, as well as the seeds, 
are borne naked on the surface of open carpels, or on the apex of 
a stem (ovary wanting). The pollen-grains are conveyed by the 
wind to the ovules, and caught by drops of mucilage, secreted by 
the micropyle. A " stigma " is wanting. The entire female prot hal- 
linin (the endosperm}, which serves for the nourishment of the 
embryo, is formed before fertilisation. The archegonia are em- 
l,nl'le<l iii the upper part if the prothallium. The pollen-grains are 
11 multicellular," i.e. there is always in their interior a distinct 
prothallium, formed by 1-3 cells, and a larger cell which gives 
rise to the pollen-tube. 

Division .5. Angiospermae. The carpels surround the ovules 
and form an entirely closed chamber (ovary), in which the ovules 
mature and ripen into seeds. The surface of a portion of the 
apex of the carpel is transformed into the "stigma," which, by a 
sticky fluid and also by hair- structures, is capable of retaining the 
pollen-grains conveyed to it by the wind, or more frequently by 
insects. The pollen-tube grows from the stigma, through the 
"conducting cellular tissue" (*/T//<"), to the ovules. The pollen- 
grains contain two cells, a vegetative and a free generative cell. 
The latter passes into the pollen-tube and there divides into two, 
one of which is the sperm-nucleus. The female prothallium, 
which is intended to serve as nutritive-tissue, is formed after 
fertilisation. Arehesronia are wantino. 

* o o 


The following- characters should be added to those already given 
on page 2 : 

The Gymnos perms comprise only trees or shrubs. The flowers 
are always unisexual and destitute of perianth (except Gnetacect) ; 
the female plant of Cycas is the only one which has no flower. 
The JIALK FLOWERS are constructed on the same type as the cones of 
the Horsetails and Club-Mosses, and are most frequently lon</ shoot* 
(Figs. 243, 258, "260 A, 267 J) bearing a number of spiral or verti- 
cillate stamens. The FEMALE FLOWERS are of a more varied struc- 
ture (see the orders). The OVULE is orthotropous (except t'odo- 
carpus which is anatropous) and projects from the carpel up- 
rightly, inverted, or horizontally; it has usually only one integument 
(compare however Taxacea?) which proceeds from the upper part 
of the nucellus, so that the embryo-sac in part is placed below 
the integuments (Figs. 251, 264). The drop of mucilage which 
catches the pollen-grains dries up and draws the pollen-grain 
through the micropyle to a space just above the nucellus the 
pollen-chamber in which the germination of the pollen-grain 

In each seed, only one of the many embryos which are formed 
proceeds to its full development. The seed is always endospermous, 
and the embryo has one, two, or a whorl of several cotyledons. 
A vigorous primary root is developed on germination. THE 
VASCULAR BUNDLES in the stem are arranged in a ring, and increase 
in thickness takes place by a closed cambium-ring which forms 
bast (phloem} externally, and wood (xyhm) internally with distinct 
annual rings, as in the Dicotyledons. Only certain of the Cycadea 1 
deviate from this arrangement. The secondary wood is very uni- 
form, as it is almost exclusively formed of tracheides with bordered 
pits, but true vessels are wanting \ this also indicates a relationship 
with the Pteridophyta (see page 202). 




The Gymnosperms are biologically lower than the Angiosperms ; 
rliev arc wind-fertilised, and without extra floral-nectaries. 

This Division embraces three classes: CYCADEJE, CONIFKRJE, and 
GNKTK.K. It is no doubt monophyletic, and has taken its origin 
from heterosporous Ferns, now extinct, most nearly related to the 
Ophioglossacese and Marattiaceas. The Cycadese appear to be the 
oldest class. The Conifera? are related to these through Ginkgo. 
The Gnetacea? are more isolated. The Division is not continued into 
the higher Flowering-plants ; it has evidently attained its highest 
development, and is now in a retrograde condition. The similarity 
which has often been pointed out between certain Coniferae and 
Lyeopodina? is only in analogous resemblances, and does not entitle 
one to suppose that there is a nearer relationship, or that the 
former take their origin from the latter. 

Class i. Cycadeae. 

The stem, is very rarely ramified. The leaves are large, pinnate, 
and arranged spirally. The flowers are diiecinns, without perianth. 

There is only one order, the Cycadaceae. In habit they 
resemble the Ferns, especially the Tree-Ferns (compare Figs. 207 
and -250). The stem is tubercular (Fig. 258), or cylindrical (Fig. 
2"><>), but not very tall (as much as about 12 metres), and very 


h-cinnlix (female plant.). The carpels are seen hanging from the top of 
stem. Three lesives with the leaflets still rolled up project almost vertically into the 
:ur, from the centre (if the crown. 

i"iivly minified. [J n Ceylon, unbranched specimens of Cycas are 
rarely met with in the wild state. The stems of C. circinalis 
occasionally branch in greenhouses.] 



The LEAVES ai-e arranged spirally, and so closely together that 
no free stem-surface is left between them, and have only a slight 
sheath (which is not amplexicaul, as in the Palms). They are com- 
pound (most frequently pinnate; in Boicenia, 
bipinnate) ; in some genera the leaves are 
rolled up in various ways, resembling the 
vernation in Ferns (Fig. 257) ; they are 
leathery and perennial. In some, stipules are 
present, as in the Marattiaceae. Groups of 
scale-leaves alternate in the majority with 
groups of foliage-leaves. 

The FLOWERS are without perianth. The 
MALE FLOWER is sometimes an enormous collec- 
tion of stamens (Fig. 258), which are flat in 
some (Cycns, Fig. 241), shield-like in others 
(Zamia, Ceratozamia) like the sporophylls in 
Horsetail (Fig. 259) ; but in all, the pollen- 
sacs are situated in large and varying numbers on the back of 
the stamens, and arranged in groups of 2-5, like the sporangia in 
the sori of the Ferns (Fig. 241 b, c). FEMALE FLOWERS are wanting 

FIG. 257. Ci/cas cir- 
cinalis. Part of a young 
leaf with circinate 

FIG. 258. A male plant of Stinger ia paradoxa (about 
5 nat. size). 

FJG. 259. Female cone 
of Zamia integrifolin (\-\ 
nat. size). The male cone 
is very similar externally. 

in Cycas, because the carpels do not terminate the apical growth of 
the stem. After a group of foliage- and of scale-leaves, a group of 
carpels is developed, which are pinnate and resemble the foliage- 
leaves, bearing on their edges a number of ovules (most fre- 

0.54 GY.MXOSPERM-E. .".-6) (Figs. 24-, 25(V); the same stem produces succes- 
sively scale-leaves, foliage-leaves, and carpels. The differentiation 
is not much more advanced than in certain Ferns (StrutUopterU, 
Blechnuin), where barren and fertile leaves of different form 
regularly alternate. The other genera have female floioers ; the 
carpels are shield-like in Zamia and Ceratozamia (Fiff. 240), 
and collected into cone-like flowers, which terminate the growth 
of the stem (Fig. 259). The number of ovules in these instances 
is two to each carpel. 

The SEKDS are large (most frequently 2-6 centimetres long) and 
plum-like ; the external layer of the testa is fleshy, while the in- 
ternal one is hard and horny. There are two systems of vascular 
bundles in the testa, one outside, the other inside the stone. The 
embryo is straight, attached to the end of the suspensor, which is 
often long, filamentous, and rolled up ; it has one or two cotyledons. 

The embryo in Ceratozamia and others is very slightly developed, at the time 
when the ripe seed is detached from the carpel ; and it is not until after sowing 
that its further development and germination proceed. This calls to miiid 
the Cryptogams, especially Sela<jinella,vihose macrospores are thrown off filled 
with endosperm ; but the oosphere is not fertilised till after the separation of the 
macrospore from the parent-plant, while in the Cycadeae fertilisation is effected 
before the separation. In Cyca* the testa may rupture, and the endosperm 
grow and become green in the light, even though no embryo has been formed. 
This also is an indication of its prothalloid nature. 

Gum-passages are present in all organs. Collateral vascular bundles, with 
spiral and scalariform tracheides, are found ; and normal thickening takes place 
by means of a catnbinm. An exception*! mode of growth is found in Cijcus and 
Enceplialartos, the cambium ceases to divide after a time and is replaced by a 
new cambium which arises in the cortical parenchyma just outside the bast, and 
which forms a new ring of xylem and phloem. This may be repeated so that a 
number of concentric rings are produced. In Ceratozamia, structures resem- 
bling corals extend from the roots iu a vertical direction and appear on the 
surface of the soil ; these are peculiar roots, in which a symbiotic Alga 
(A null, run) is found. 

The Cycadeaa were formerly (from the Coal period to the Later Cretaceous) 
far more numerous than at the present day. They appear to have been most 
numerous in the Trias and Jurassic. The remnant (75 species) which have 
persisted to the present time are found in all tropical countries. Cycas (Trop. 
and Sub-trop., Eastern Hemisphere) ; Dioou (Mexi -ol : Macrozamia (Australia) ; 
K.v/,/,,,/,ir/.is (Trop. and S. Africa) ; Stangeria (Fig. 258, Sub-tr p. South and 
Eisl Uiir.ii: l',,ir,'nia (Trop. Australia); Ceratozamia (Mexico, New Granada, 
\\Y<t.Tn Brazil) ; Microcijcas (Cuba) ; Zamia (Trop. and Sub-trop. N. America.) 
Sago is made from the starch-containing pith of Cycas revuluta and 
11-,-iiiiili.t. The leaves are often used at funerals and church festivals, under 
the name of " palm-branches." 


Class 2. Coniferae (Pine-trees). 

The stem branches freely. The leaves are entire, relatively 
small, linear or reduced to scales. The flowers are without peri- 
anth. The ovules naked. It is seldom that the female flower is 
reduced to only one carpel. 

Whilst the Cycadeae principally resemble the Ferns, the Conifers 
partly resemble the Lycopods, and partly the Equisetums the 
former especially in the needle- or scale-like, leathery, simple, and 
often perennial leaves (" evergreen plants "), which never posaegs 
stipules (Figs. 263, 270, 272). G ink-go deviates from this, being 
no doubt the oldest, and the Conifer which stands nearest to the 
Cycadea? (Fig. 260). The resemblance to the Equisetums is 
especially owing to the fact that the stem ramifies abundantly, and 
often very regularly, forming a pyramid with verticillate branches. 
In addition to the foliage-leaves, scale-leaves (bud-scales) are 
present in the majority of species. 

The FLOWERS are monoecious or more rarely dioecious. Perianth 
is wanting. The stamens of the catkin-like male flowers (Fig. 267, 
J) are of different forms, but as a rule more or less shield-like. 
As in the Cycadeae, the pollen-sacs are in all cases situated on the 
underside. There are, as a rule, two pollen-sacs (the Abietaceae, 
Fig. 267), or 3-5, (the Cupressaceaa and Taxaceaa, Fig. 243) ; a 
few have more, e.g. Araucaria (Fig. 242) ; they dehisce by clefts. 

If, in commencing our consideration of the female flower, we 
begin with that of Giithgo, we shall observe in the corner of a scale- 
or foliage-leaf a small flower, which consists of two carpels, each 
bearing one ovule, and reduced almost to the ovule itself (Fig. 2(50 
C, D). The flower in Podocarpns is still further reduced, viz. to a 
single carpel with one ovule, which is anatropous and has two in- 
teguments. This ovule is situated in the axil of a cover-scale (c, in 
Fig. 262 D), and several female flowers of this description are col- 
lected in a small cone, the stalk and bracts of which become fleshy 
( Fig. 262 C). The external integument also becomes fleshy (an aril). 
Dacrydium, which is clearly related to Podocarpus, has an external 
integument which developes more independently as a fleshy aril 
(Fig. 262 B, I? 1 ). Microcachrys also is clearly allied to these : 
the bracts are more fleshy, and the ovule (i.e. the female flower) 
is protruded beyond the bract (Fig. 262 A, A 1 ). Taxus stands 
in a more isolated position : a flower which has been reduced to 
an ovule is situated, in this instance, on the apex of a secondary 


branch which is studded with floral-leaves (Figs. 263, 264); an ex- 
it-mill integument is developed on all sides and surrounds the seed 
as a scarlet aril. According to this conception the aril corresponds 
in an eternal integument, and the Taxoideae thus possess a partly 
dichlainvdeons ovule. Only Ginkgo and Cephalotaxus appear to 
deviate from this, as in these there is only one integument (unless 
the small outgrowth indicated by ar, in Fig. 260 D, really is a 
rudimentary, external integument) ; in CYCAPEJE, to which Gink go 
is most closely related, there is likewise only one integument. But 
in these genera the testa is differentiated into two layers, and the 
seed resembles a drupe ; like the Cycadese there is an external 
fleshy covering and an internal hard one, and these two layers 
mav probably be considered homologous with the two integuments. 
This theory is also borne out by the arrangement of the vascular 
bundles in Ceplilot<t,fns and Podocarpus, which present the xylem 
in the fleshy external layer to the outside of the testa, which is 
therefore the upper side of the integument (Celakovsky). 

The coalescence of the integuments into one is only slight in 
Torreya, more pronounced in Podocarpus and strongest in Cephalo- 
taxus and Ginkyo. Celakovsky terms these ovules " holochlamy- 

If we pass from these to the order PIXOIDE.E, we find the female 
flowers collected into catkin-like cones, which have been considered 
from various points of view to be sometimes single flowers, at 
t>t her times compound inflorescences. The structure in ABIETACE.E 
is as follows : a number of spirally arranged, scale-like leaves, 
cover-scales (Figs. 267, 268), are situated on a long axis. In the 
axil of each cover- scale a larger leaf-like projection, the ovuliferous 
scale, is borne, which turns the upper side towards its cover-scale 
(which is shown by the fact that the wood of its vascular 
bundles is turned downwards and towards the wood in the bundles 
of ihe cover-scale : Fig. 269). Two ovules, with micropyles turned 
towards the central axis, and with apparently only one integument 
( Fig. 268), are situated on the dorsal side of each ovuliferous 
scale, i.e. the side turned away from the cover-scale. The ovuli- 
trnms scales grow after fertilisation, into the woody or leathery 
"cone-scales," which are usually much larger than the cover-scales. 
Tin.-, ovuliferous scale with its axis may, according to Celakovsky, 
! considered as a dwarf-branch which is situated in the axil of 
tin- cover-scale, and bears two ovules (in the same way as in 
Oinlcgo, one long-stalked flower, reduced to two ovules, is situated 



in the axil of a leaf), and in this case the external integument of iln 1 
ovules is expanded into leaf -like bodies, which have united to form 
one " symphyllodium " (pvuliferous scale) which is inverted so that 
its dorsal side is turned upwards and bears the nucellus and 
the other integument (" hemichlamydeous " ovules). The carpel 
itself is therefore in this instance extremely reduced. The keel, 
or (in Pw?f.s-) " macro" (Fig. 268 5), which is foand in several 
genera, represents then a third carpel, which is sterile. In the 
other orders of the Pinoidese the cover-scales and ovuliferous scales 
grow more and more together and finally form one structure, which 
also is termed a " cone-scale," although from its development it 
cannot be homologous with the cone-scales of the Abietacea?. 
This connation is least in the TAXODIACEJI and ARAUCARIACE^ and 
may be traced on the upper surface of the " cone-scale " by the 
presence of a stronger or slighter ridge or pad, the free portion 
of the ovuliferous scale (Figs. 256, 266, 269). It is most strongly 
pronounced in the CUPRESSACEJE, in which the two scales form one 
single structure, the cone-scale (Fig. 274). The vascular bundles 
in the under portion corresponding to the cover-scale, have the 
xylem towards the upper side as usual in leaves, whilst the bundles 
present in the upper side of the cone-scale, which thus represents 
the ovuliferons scale, turn their xylem downwards. The hemi- 
chlamydeous ovules are then situated on the upper side of this 
cone-scale. According to this theory the CUPRESSACEJ: appear to be 
the youngest type, a view which corresponds with their vegetative 
structure. If there is only one ovule in these orders as in Ayathi* 
(Fig. 265) and Araucaria, then the flower is reduced to a single 
carpel and one ovule, as in the case of Dacrydium and Microcachrys. 
If two or more ovules are present, then the same number of carpels 
may be supposed to exist, the external integuments of their 
ovules being developed into leaf-like structures which collaterally 
coalesce to form a " symphyllodium," or are suppressed. 

According to this theory, which is based on the researches of 
Celakovsky, the female flowers of the Coniferae may be classed 
thus : 

1. In all cases situated in the axil of a bract and collected into 
cones, with numerous flowers or with few or one flower. In Ginkgo 
only, are they situated in the axil of foliage- or scale-leaves. 

2. It is only in Taxus that bracteoles are pi'esent. 

3. They are formed only from rudimentary carpels, in which the 
stem takes no part. 

w. B. s 


4. The number of carpels in each flower varies from one to 
many, most frequently three, of which the central one remains 


5. Each carpel bears only one ovule. The flower which is 
formed of only one carpel appears to consist of only one ovule. 

'-. The ovule has in Taxaceee either a double integument 
(Podocarpese, Taxea>), of which the external is the "aril," or, as 
in the Cycadea?, a single one, which is homologous with the two 
united together. 

7. The external integument in the Pinoidea? is expanded to 
form a leaf-like structure the ovuliferous scale and bears on its 
dorsal side the ovules, which are thus only provided with one, and 
that the inner, integument. 

This later interpretation of the female cones in the Coniferae is more pro- 
bably correct than the older ones ; that, however, which appeared in the former 
issues of this book, may also be stated. It was to the effect that each catkin- 
like female cone is in reality a single flower ; the cone-scales in the Cupressaceae 
were single leaves, namely carpels, which bore the ovules on the side which is 
turned upwards ; the division into two parts which makes its appearance in the 
other orders, and becomes most prominent in the Abietacerc, was compared 
with the division of a leaf into a barren and a fertile portion, which is found 
especially in Ophioglossaceas and Marsiliacese, or with the ligule in Isoetes. 

POLLINATION is accomplished by means of the wind. At the 
period of pollination the leaves are always so widely separated 
from one another, that the ovules can catch the pollen-grains 
carried to them by the wind; this is often effected by the mucila- 
ginous drops which appear at the micropyle, and by the evapo- 
i at inn of which the pollen-grains are brought in contact with 
tin- nucellus. The entire cone grows considerably as soon as 
fertilisation has taken place, and the cone-scales in Pinoidea? 
cloSe together so that the seeds while maturing are enclosed, and 
it i> not until the seeds are ready for distribution that the cone- 
scales again become separated. In the Pinoidea-, the fully deve- 
loped ovuliferous scales are hard and woody ; and in this condition 
tin- collection of female flowers is termed a cone. In the Taxoidece, 
true cones are the exception. 2-15 cotyledons are present, ar- 
i-an-vil in a whorl. 

''In- '-liararteristic feature of this class is the abundance of 
which is to be found in isolated cells (especially in the 

CONIFER*. 259 

cortex), partly in intercellular glands or passages (both in the 
cortex and wood). Taxus is the only genus which has no resin. 

There are about 350 species, mostly from the Northern Temperate zone 
(especially North America and Siberia), where they grow gregariously and form 
the most northern forests. The Juniper, Scotch Fir, and Yew are natives of 
Great Britain. 

This class may be divided into two families : 

1. Taxoideae. The ovules have either one integument, the 
external part of which is fleshy, and the internal hard and stone- 
like ; or two integuments, of which the external is the fleshy and 
coloured "aril." "Ovuliferous scales" are wanting. The cones 
are never woody, but are generally succulent, the bracts become 
fleshy, or cones usually are not developed. The seeds project more 
or less freely beyond the bracts. 

2. Pinoideae. The ovules have two integuments, the external 
one of which is leaf-like and becomes developed as the " ovuli- 
ferous scale " ; if there are several of these in each flower they 
unite and form a " symphyllodium." This may remain free or 
unite with the bract. The cones are most frequently woody, 
rarely succulent. The seeds are hidden among the cone-scales. 

Family 1. Taxoideae. 

This family, considered to be most nearly related to the Cycadece, 
also made its appearance at a very early period. There is only 
one order. 

Order. Taxaceae. The characters have been given above. 

A. CEPHALOTAXEJ; is the oldest group, presumably the connect- 
ing link between the Cycadece and the other Conifera?. The flower 
consists normally of two ovules. Aril wanting. One integument. 
Seeds drupaceous. The flowers in Ginkgo bilola (Salisburia) 
sire situated in the axil of foliage- or scale-leaves. The stamens 
bear only two pollen-sacs (Fig. 260 A). The female flower has 
two ovules, placed together at the end of a long, bare axis (Fig. 
260 (7). Round the base of the ovule a small collar (ar, in Fig. 
260 .D) is found, which may probably be considered homologous 
with the collar-like outgrowth which surrounds the base of the 
Cycas-ovn\e. The seed resembles a Plum, and has a fleshy external 
coat, surrounding a hard internal layer. The embryo is developed 
after the seed has fallen off. The Ginkgo-tree has long-stalked, 
fan-shaped leaves, more or less indented, with dichotomous veins 



resembling certain Ferns the Adiantums. It is a native of East 
A. -in, and the only surviving species of a genus which in earlier 

FIG. 260. G in l:go (nat. size) : A a branch with a small flowering dwarf-branch (male 
flower); B a leaf ; C a flower with two ovules ,- Da ripe seed; ar collar. 

times was very rich in species, and distributed over the entire 

Northern. Hemisphere. Ceplialotaxns (Eastern Asia) is related to it. 
13. PODOCARPO:. The female flower is reduced to one ovule, 

placed in the axil of a bract, or a little forward upon it. The ovule 

has an aril (2 integuments). Phyllocladus 
(Fig. 261), from New Zealand and Tasmania, 
has obtained its name from its flat, leaf-likf 
branches, the leaves proper being scale-like 
(/). The ovules stand erect in the axil of 
the scale-like leaves (c), and several are 
collected at the end of short branches. 
Microcachrys tetragona (Tasmania) has a 
small female catkin with several spirally- 
placed, fleshy bracts, at the end of which 
the inward and downward turned ovule is 
attached (Fig. 262 A, A'). The ripe cones 
are red, succulent, and resemble Straw- 
berries. In Dacrydium (Tasmania, New 
- branch C '"'!tu Zealand > Malaysia) the female cone has most 

female flowers (nat. size). frequently only 1-2 (-6) bracts, which re- 



serable the vegetative leaves ; they have also a fleshy aril (Fig. 
262 1>, !>'). Podocarpus (40 species, East Asia, S. Temp.) ; the 
bracts of the female flowers become fleshy, and unite together ; 
only 1 or 2 are of use in supporting the flowers. The ovules 


FIG. 262. .4 Microcaclirys : female cone (J). A' A single carpel with its ovule. H 
Dacrydium : branch with female flower (~). B' The flower; cp the bract; ar the aril; ov 
ovule. C Podocarpus : female flower with 2 ovules. D Another female flower with 1 ovule, 
in longitudinal section. 

project high above the apex of the bract, and are anatropous, the 
micropyle being turned downwards (Fig. 262 C, -D). An aril 
commences to develope in the flowering period as an external 
coating, and later on it becomes fleshy and coloured. 

C. TAXE.E. The female flower is reduced to one ovule, which is 
situated terminally on an axis which bears 2-3 pairs of opposite, 
scale-like bracteoles ; on this account the Taxere form a very 
isolated group among the Coniferse. 
-Taxus (T. baccata, the Yew-tree). 
Dioecious. The female flower con- 
sists of only one ovule, placed at the 
end of a short secondary branch 
(Fig. 264), which is studded with 
scale-like leaves. The aril when 
ripe is thick, fleshy, and scarlet 
(sometimes yellow), and only 
loosely envelopes the seed (Fig. 
263). The leaves are scattered, 

. FIG. 263. Taxus linccata .- branch 

flat, linear, and pointed (Fig. 263, with two ripe seeds (nat. size). 


264). The short male flowers have 5-8 pollen-sacs, pendent from 
the stamens, and are surrounded at their bases by scale-like 
bracteoles (Fig. 243). Torreya (4 species, N. America and Japan) 
is closely allied to Taxus. The aril ultimately fuses with the 
woody inner integument, and hence the ovule becomes drupaceous, 
us in Cephalotaxese. 


FIG. 261. Tn.i'us Iciccttfa: A shoot of 'J'n.nm with female flowers at the time when the 
ovules are ready for pollination. B Leaf with flower in its axil (nat. size). C Longitudinal 
median section through a female shoot ; v growing point of primary shoot ; a commence- 
ment of aril; i integument; nnucellus; m micropyle. 

USES. Tn.rux liat-cata is usually planted iu gardens, especially in hedges. 
Its wood is very hard and is used for wood-carving. The shoots are poisonous, 
but not the aril, \vhich is often eaten by children and by birds. 

Family 2. Pinoideae. 

The four orders differ from one another partly in the arrange- 
ment of the leaves (Cupressacece have opposite or verticillately 



placed leaves, flowers, and inflorescences ; in the others they are 
placed spirally), but chiefly in the greater or less degree of union 
which takes place between the female flower (the leaf-like " sym- 
phyllodiurn") and its supporting cover-scale, and in the position of 
the ovules (the micropyle being turned upwards or downwards). 
The "cone-scales" in Abietacece are formed by " symphyllodia " 
alone, in the others by their union with the cover-scale. 

Order 1. Araucariaceae. This order most frequently has 
solitary ovules, turned doicmvards and attached to the centre of the 
cone-scales. In Agathis (Dammara) the arrangement is the most 
simple, a winged seed (Fig. 265), which hangs freely downwards, 
being borne in the centre of the undivided cone-scale. In Arau- 
caria, the stamens with the free, pendulous pollen-sacs have been 
represented in Fig. 242 ; the ovuliferous scale is united for nearly 


PIG. 265. Agathis (Dam.rn.ara) au&tralis. Cone-scale with the seed. A Longitudinal 
section ; A' from within ; fu,fv' vascular bundles ; v wing. 

FIG. 266. Cunninyhamia sinensis. Cone-scale with three ovules, interior view: d cover- 
scale ; / ovuliferous scale. 

its whole length with the bract, and projects from its apex in the 
shape of a sheath-like, dentate scale, resembling the ligule in 
Isoiites, and may therefore be termed a " lignle." Araucaria (S. 
America, Australia) has often rather broad leaves (A. brasiliensis). 
The ovuliferous scale in Cnnninghamia is more distinct, and 
stretches transversely over the entire cover-scale ; it bears three 
inverted ovules (Fig. 266) (Eastern Asia). 

Dammara -resin, which is used for varnish, is obtaiuel from Agatliis 
(Dammara) species (New Zealand, Philipj ine Islands). 

Order 2. Abietaceae (Pine and Fir Trees). The leaves 
ara spirally arranged and needle-like. The flowers are monoecious. 



The male flowers are long, and catkin-like, with numerous stamens, 
each bearing 1 two oblong pollen-sacs. The pollen-grains are most 
frequently tri-lobed, having two bladder-like appendages, formed 
its outgrowths of the exospore, to assist in their distribution by the 
wind (Fig. 267 N). The bracts are arranged spirally. The union 
between the bract and the ovuliferous scale, which is found in the 
preceding order, is not in this instance so complete ; these scales 
make their appearance as two free parts, and are attached only 
at their bases (Fig. 268) ; the lower portion, that is the cover- 
scale, in most instances remains quite small (Fir, Red Pine, and 
others), it is only in the "Noble Pine " (Abies) and Pseudotsm/u 

FIG. 267. A-G Pseudotsuga douglasii : A cone, B cone-scale, with the inner side 
turned forward ; the points of the cover-scale are seen behind it ; C-G transitions from 
the acicular leaf to the cover scale, from the base of a $ cone. H Finns montana. Young 
ovuliferous scale, with the inner side turned forward; the ovules are now in the stage for 
pollination. J-M Allies alba: J male cone ; b bud-scale; o anthers; KLM individual 
anthers. Pinus wotitana : N pollen-grain; the two lateral expansions are the air- 
Madders; in the upper part of the interior of the grain a vegetative cell may be seen, 
and in the centre the large cell-nucleus. 

ii, that it attains a greater length than the ovuliferous 
scale (Fig. 267, B-G). On the other hand the upper part, the 
ovuliferous scale (the vascular bundles of which have the bast 
turned upwards), grows strongly and elongates, especially after 
fertilisation, becoming woody or leathery ; it is commonly termed 
the " cone-scale ," but is in reality only homologous with a part of 



the ; ' cone-scale" in the other order of Pinoidese. On the side 
of the ovuliferous scale, turned towards the axis, are situated two 
ovules with micropyles directed inwards. The seeds are most 
frequently provided with a false wing (a tissue-like part of the 
surface of the ovuliferous scale). Cotyledons, more tluut '2, 
verticillate. Fertilisation, does not take place until some time after 
pollination. In Pinus, for instance, the pollen-tube only penetrates 
the nucellus for a short distance during the year of pollination, and 
then ceases its further growth, fertilisation not taking place until 
after the middle of the next year; whilst the seeds ripen about a 


FIG. 263. .4 Abies : c the cover-scale ; s ovuliferous scale, or " cone-scale " ; sic ovules in 
a .young condition. 1> Pinus : ovuliferous scale with two ovules (s) ; m the tvro-lobed 
micropyle; c " mucro " ; b the cover-scale behind. C Abies : ripe "cone-scale" with two 
seeds (su); /winy of seed. 

year and a half after pollination. In the Larch and others, the 
seeds are mature in the autumn succeeding pollination. 

Abies (Fir). The leaves are often (e.g. Ab. pectinata) displaced 
into 2 rows, flat and indented at the apex, with 2 white (wax- 
covered) lines on the under surface, in which the stomata are situ- 
ated. The leaf-scars are nearly circular and do not project. The 
cones are erect. The cover-scales and the ovuliferous scales separate 
from the axis, to which they remain attached in other genera. 
Tmiga has leaves like Abies, but by the slightly projecting leaf-scars, and cones 
with persistent scales, it forms the transition to Picea. Pseudotsuga has leaves 
similar to those of Abies and persistent carpels as in Picea, but the cover-scales 
grow as in Abies and project beyond the ovuliferous scales (P. dotiglasii, Fig. 
267). These two genera are considered as sub-genera of Abies. Picea. The 
leaves project on all sides, square and pointed ; the leaf-scars are 


rhombic, on projecting leaf-cushions. The cones are pendulous. 
The cover-scales are much shorter than the leathery, persisting 
ovuliferous scales. The genus Larix (Larch) differs from all the 
others in having deciduous leaves (the three preceding have 
leaves which persist for eleven to twelve years). It has long- 
brandies with linear foliage-leaves and short, thick, perennial 
dwarf -brandies, which each year form a new rosette of foliage- 
leaves, similar to those on the long-branches. The male flowers 
and the erect cones resemble those of Picea, and are borne on 
dwarf-branches. Cedrus (Cedar) resembles Larix to some extent, 
but has persistent leaves (C. libani, C. deodara). Pinus (Pine) has 
long-branches and dwarf-branches. The leaves of the long- 
branches are scale-like and not green; the dwarf-branches have 
very limited growth, and persist for three years; they arise in 
the axils of the scales borne on the long-branches of the self- 
same year, and each bears 2-5 foliage-leaves, they are also sui 1 - 
rounded at the base by a number of membranous bud-scales. 
The cone-scales have a thick, rhomboid extremity (the ''shield"). 

The buds which develope into long-branches arise at the apex of other long- 
branches, and being very close together, form false whorls. The female cones 
occupy the position of long-branches, and take about two years for their develop- 
ment. The male flowers arise close together, and form a spike-like inflorescence 
at the base of a long-branch of the same year. The male flowers occupy the 
position of dwarf-branches, so that a female cone may be considered to be a 
modified long-branch, and a male cone a modified dwarf-branch. The main 
axis of the seedling hss needle-like leaves, similar to those found on the older 
parts, and on dwarf-branches ; it is not until some time later that the dwarf- 
branches are developed and the permanent arrangement attained. 

USES. Several species are commonly cultivated in this country, partly on 
heaths and moors, and partly in plantations and as ornamental trees, such 
as Mountain Pine (Finns montana, Cen. Ear.); Austrian Pine (P. larieio, 
Eur.) ; Scotch Fir (P. silvestris, Eur.) ; Weymouth Pine (P. strobtts, N. Am.) ; 
common Ked Pine (Picea excelsa, Cen. and N. Eur.) ; White Pine (P. alba, N. 
Am.) ; Abies pectiuata (Common Fir, S. and Cen. Eur); A. nonbnanni ina 
(Crimea, Caucasus); A. balaamea (N. Am.); Tsnya camtdcmis (N. Am.); 
Pneudotsuga douyladi (W.N. Am.) ; Larch (Larix europcca, Alps Carpathians) ; 
L. sibirica {N.E. Russia, Siberia). The wood of many species, especially Pine, 
on accoui.t of its lightness and because it is so easily wo ked, is very well 
adapted for many useful purposes. The wood of the Yew-tree is very hard 
and is used for ornamental turning. Resin and Turpentine (i.e. Resin with 
essential oils, the name being derived from the Terebinth-tree, from which 
formerly a similar material was obtained) are extracted from Finns larieio and 
P. pinaster. Oil of Turpentine is obtained by distillation of turpentine with 
water; Tar by dry distillation of Pine-wood. Canada-balsam is from North 



American Abies-species (A. bahamea and Fraseri}. The officinal Turpentine 
is mainly obtained from Finns pinaster (South of France), P. t<mhi, aitttralis, 
strobus (Weymouth Pine) and other North American species ; more recently 
also from P. silvestris (Scotch Fir), maritima, laricio, Picea excelsa, and others; 
Venetian Turpentine, from Larch (S. Eur.) Amber is resin from a Tertiary 
plant (Pityoxylon succinifenun), closely related to the Pine, which grew especially 
in the countries round the South-East coast of the Baltic. Pi mis piiiea (the 
Pine, S. Eur.) has edible seeds and also P. cembra (in Ceu. Eur. and Siberia). 

Order 3. Taxodiacese. The vegetativ 
are arranged spirally. The ovules (2-9) 
are situated either at the base of the 
ovuliferous scales, in which case they 
are erect ; or at their centre, when they 
are generally more or less inverted. The 
ovuliferous scale is more or less united 
with the cover-scale, and projects be- 
yond the surface of the cone-scale, like a 
comb (Fig. 269). The vascular bundles, 
which extend into the cover-scale, have 
the usual leaf-arrangement, viz. the 
wood placed above the bast ; while those 
bundles which enter the ovuliferous 
scale have this arrangement of the 
bundles reversed. 

e leaves and cone-scales 

FIG. 269. Cryptomeria ju- 
ponica. Portion of longitudinal 
section through female flower. 
d cover-scale ; / ovuliferous 
scale ; ou ovules ; J'v and fo' 
vascular bundles; the xylem is 
indicated by a wavy line, and 
the phloem by a straight line. 

Taxodium ilistichiim (the North American "Swamp Cypress") has annual 
dwarf-branches, with distichous leaves, and cone-like " pueumathodia." In the 
Tertiary period it was very common in the Polar regions. Sequoia (Welling- 
ton ia) yigantea is the famous Californian Giant-Fir, or Mammoth-Tree, which 
attains a height of BOO feet, a diameter of 36 feet, and is said to live for 1,500 
years. Cryptomeria japonica (Japan, China) has the least aduate ovuliferous 
scales ; Glyptottrobus (China) ; Arthrotaxis (Tasmania) ; Sciadopitys verticillata 
(the only species in Japan) has, like Phius, scale-like leaves on the long-branches, 
of which those which are situated at the apex of the annual shoots support 
" double needles," i e. dwarf -branches similar to the two-leaved dwarf-branches 
in Pimm, but without bud-scales, and with the two leaves fused together at the 
edges into one needle, which turns its upper surface away from the long-branch. 

Order 4. Cupressaceae (Cypresses). The leaves are opposite 
or verticiUate, sometimes acicular, but most frequently scale-like 
(Fig. 270). In the species with scale-like leaves, the seedlings 
often commence with acicular leaves (Fig. 272), and branches- 
are sometimes found on the older plants which revert to this 
form, seeming to indicate that the acicular leaf was the ori- 



ginal form (atavism). The so-called " Eetinospora " species are 
seedling-forms of Biota, Thuja, Chamcecyparis, which have been 
propagated by cuttings, and retain the seedling-form. The flowers 
are monoecious or dioecious. The male flowers are short, and 
have shield-like stamens, bearing most frequently several pollen- 
sacs. The cover-scales and ovuliferous scales are entirely fused 
together and form undivided cone-scales, opposite or u-horled; the 
orttliftTuus scales have slight projections near the base on which 
l-2-several erect ovules are developed (Fig. 274). Most frequently 
2 cotyledons. Evergreen trees and shrubs. 

FIG. 270. 

FIG. 271. Portion of a branch 
of Thuja orientalisl (magnified). 
The leaf at the base on the right 
has a branch in its axil. 

FIG. 272. Seedling of 
Thuja occidentalis. The 
branch (g) is borne in 
the axil of the leaf s. 

Juniperus (Juniper). Dioecious. The cone-scales become fleshy 
and fuse together to form most frequently a 1-3 seeded 
"berry-cone." J. communis (Common Juniper) has acicular leaves, borne in 
whorls of three, and the " berry-cone " is formed by a trimerous whorl of coue- 
scales (Fig. 273). J. sabina and J. virginiana have " berry-cones" formed from 
several dimerous whorls of cone-scales; the leaves are connate and opposite, 
needle- and scale-like leaves are found on the same plant. 

(Cypress). Monoecious. The cones are spherical ; the 
cone-scales shield-like, generally five-cornered and woody (Fig. 
2~n), each having many seeds. The leaves are scale-like. Thuja. 



Monoecious. Cones oblong. The cone-scales are dry, as in the 
Cypress, but leathery and imbricate, and not shield-like ; each 
cone-scale bears 2-3 seeds. The leaves are most frequently 
dimorphic; those leaves which are situated on the edges of the 
flat branches are compressed, and only these bear buds, which 
are developed with great regularity, generally alternately, on both 
sides of the branch ; those which are situated on the flattened 
surfaces are pressed flat and broad, and never bear branches 
(Fig. 271). Along the central line of each leaf there is a resin- 
canal (Fig. 271 ). Chanxecyparis, Callitris, Libocedru*, Thujopsis (1 species : 
T. dolabrata ; in Japan). 



FIG. 273. Branch of Juniper with 
" berry-cones." 

FIG. 27i. Cupressus lau'soniana. Loii'ji- 
tudinal section through female cone. Two 
ovules (OB) are bisected; / ovuliferous scales. 

OFFICINAL. Junipenis sahiiia from Central and South of Europe (the young 
branches yield an essential oil). The wood of J. communis is used in the 
production of an essential oil, and J. oxycedrus in the production of empy- 
reuinatic oil. The ' berry-cone " of J. communis is officinal, and is also need 
for gin. The wood of J . virgimana (N, Am.) is known as red cedar, and is 
used for lead-pencils. Sandarack resin is obtained from Callitris quadrii-iilrii 
(N.W. Africa). 

THE FOLLOWING ARE CULTIVATED is GABDENS : Thuja occiclcittalif! (Arbor vitas) 
(>J. Am.), and orientalis (China, Japan) ; Juiiipeni* saldna and virginiaim ; 
Tltnjo2)sis dolalrata (Japan); Cupressus lawsoniana (California), C. sempervirens 
(S. Eur., W. Asia), and other species, are grown especially in conservatories, 
and in Southern Europe particularly in cemeteries. The Retinospora species 
which are so often planted, do not belong to an independent genus, but are 
obtained from cuttings, taken from seedling-plants with acicular leaves (see page 



Class in. Gneteae. 

Tliis class, independent of extinct forms, comprises the most 
highly developed of the Grymnosperms, partly from the circum- 
stance that a perianth of 2-4 members encloses the terminally 












GNETE-ffi. 271 

placed ovule, which is provided with one, or (in Gnetiini) two, 
integuments, and partly owing to the fact that the wood has true 
vessels. There is only one order. 

Order. Gnetaceae. The three known genera differ very much in appear- 
ance. Welwitwhia mirabilis (from the deserts of South Western Africa) is the 
oldest (?) genus now living. It resembles a giant radish, in that the hypocotyl 
is the only part of the main axis of the stem which becomes developed. It 
attains a circumference of upwards of four metres with a length of A-$ of a 
metre. It bears only two oblong, leathery leaves (Fig. 275) which are torn into 
segments at the apex and lie on the surface of the soil ; these are the two first 
foliage-leaves which succeed the cotyledons, and they are remarkable for their 
enormous length (upwards of two metres) as well as for their long duration, 
living as long as the plant itself. In their axils are situated the 4-rowed, spike- 
like male and scarlet-coloured female cones, upon dichotomous branches. The 
perianth consists in the $ of '2 alternating pairs of leaves, the inner ones of which 
are slightly united. The androecium likewise consists of "2 whorls : the external 
(transverse) with 2, the internal with 4 stamens ; the lower halves of the 6 
filaments uniting to form a cup. Each of the terminal anthers corresponds to 
a sorus of 3 sporangia, the sporangia being fused together, and opening at the top 
by one three-rayed cleft. In the centre of the $ -flower there is a sterile 
ovule. In the $ -flower a perianth of two connate leaves is present. Epliedra 
(desert plants, especially in the Mediterranean and W. Asia) at first sight re- 
sembles an Equisetum;the stems are thin, long- jointed, and the leaves opposite, 
small, and united into a bidentate sheath ; $ -perianth of two connate leaves 
(median leaves) ; 2-8 stamens united into a column. Each anther is formed 
of 2 sporangia (is bilocular). $ mainly, as in Weltcitschia. The seeds are sur- 
rounded by the perianth which finally becomes red and fleshy. There are 30 
species. Onetum has opposite, lanceolate, pinnately-veined, leathery leaves. 
They are mostly climbers (Lianas) from Tropical Asia and America. The $ - 
flowers have a tubular perianth, (formed from two median leaves) which 
surrounds a centrally-placed filament, bearing 2 anthers. In the $ -flower 
there is a similar perianth, surrounding an ovule provided with 2 integuments. 
The perianth becomes fleshy and envelops the hard seed. 20 species. 

From the circumstance of Welvaitschia having $ flowers which, bssides 
stamens, possess also a rudiment of an ovule, Celakovsky draws the inference that 
the earliest Gymnosperms had hermaphrodite flowers which from this structure 
became differentiated entirely into $ - and $ -flowers, with the exception of 
IVelivitschia only, in which this differentiation was only carried out iu the $ - 
flower. This theory has so far been scarcely proved. 

Fossil Gymnosperms. 

The earliest continental plants which are known belong to the CORDAITACE.&, 
a group of plants which existed as early as the Silurian period ; they were 
Gymnosperms, but it has not yet been determined whether they were Cycads or 
Conifers. The CYCADS, even in the Coal period, were scarce ; they attained their 
fullest development in Jurassic and Cretaceous periods, during which they were 
rich in species and genera, and extended as far as the Polar regions. In addition 

to these, Taxacea?, Abietaceas, and Taxodiacea? appeared in the Carboniferous 
period. The TAXACE.E appear to have attained their culmination in the 
Jurassic and Cretaceous periods ; (linkyo appears in the Rhaetic ; Torreya, in the 
Cretaceous ; 7' and Podocarpiis in the Tertiary periods. The ABIETACE.K 
also appear in the Carboniferous ; Finns was first known with certainty in the 
English Weald and in the Cretaceous ; almost all other contemporary genera are 
represented in this latter period. The ARAUCARIACE.E first appear, with certainty, 
in the Jurassic. The TAXODIACE.E may be traced back as far as the Carboni- 
ferous (?) ; Sequoia is first found in the lowest Cretaceous, at tbat period it 
spread throughout the entire Arctic zone, and being represented by a large 
number of species, formed an essential part of the forest vegetation. Sequoia 
played a similar part in the Tertiary period. The CUPRESSACE^: are first 
known with certainty in the Jurassic, but they appeared more frequently and 
numerously in the Tertiary period, in which most of the present living genera 
were to be found. The GNETACE^E, according to a theory advanced by Renault 
were represented in the Coal period by the genus Stephanospermum, which had 
four ovules enclosed by an envelope. 


See pages 3 and 224. To this Division belong the majority 
of the Flowering - plants. They are divided into two parallel 
classes, the Monocotyledons and the Dicotyledons, which differ 
from each other not only in the number of cotyledons, which, with 
a few exceptions, is one in the former, two in the latter, but also 
in the internal structure of the stem, the venation of the leaves, 
the number of the parts of the flower, etc. Assuming that these two 
classes have sprung from a common origin, it is amongst the Helobieae in the 
first, and amongst the Polycarpicte in the second class that we might expect 
to find closely allied forms, which might reasonably be supposed to have varied 
less fr-im this original type. As for the rest, they seem to stand quite parallel, 
without exhibiting any close relationship. It is scarcely proved that the 
Monocotyledons are the older class. 

[Our knowledge of the forms included under the Angiosperms 
has recently been considerably increased by Treub (Ann. d. Jar. 
Hot. d. Bidtenzorg, 1891), who has shown that the Casuarinas differ 
in many important points from the typical Angiosperms. Among 
other characters the pollen-tube is found to enter the ovule near 
the chalaza and therefore at the opposite end to the micropyle, 
and Treub therefore suggests that these plants should be placed 
in a subdivision termed Chalazogams. 

According to this view the principal divisions of the Angio- 
sperms would be represeiited thus : 


Sub-division. Sub-division. 


Class. Classes. 

Chalazogames. Monocotyledones, Dicotyledones. 

More recently Nawaschin (Bull. Acad. Imp. Sci. St. Peterxb., 
ser. iii., xxxv.) has shown that Betula, and Miss Benson (Trans. 
Linn. Soc., 1894) that Alnus, Corylns, and Carpinus also belong to 
the Chalazogams. 

w. B. 273 T 

274 ANGIOSPEBM.ffi. 

Our knowledge, however, is still so incomplete that one would 
hesitate to accord the full systematic value which Dr. Treub 
attaches to his discovery until the limits of the Chalazogamic 
group are better denned ; and it would hardly be justifiable to 
include the Casuarinas and the above-noted genera in one 

Class i. Monocotyledones. 

The embryo lias only one cotyledon; the leaves are as a rule scattered, 
with parallel renal ion ; the vascular bundles of the stem are closed, 
there is no increase of thickness. The flower is typically constructed of 
five %-merous u'horls, placed alternately. 

THE EMBRYO is generally small in proportion to the abundant 
endosperm (exceptions, see Helobiece), and its single cotyledon is 
often sheath-like, and very large. On the germination of the 
seed either the entire cotyledon, or its apex only, most generally 
remains in the seed and absorbs the nutritive-tissue, while the 
lower portion elongates and pushes out the plumule and radicle, 
which then proceed with their further growth. The primary root 
in most cases soon ceases to grow, but at the same time, however, 
numerous lateral roots break out from the stem, and become as 
vigorous as the primary root, or even more so. Increase in 
thickness does not take place in these roots ; they branch very 
little or not at all, and generally die after a longer or shorter time. 

THE STEM is frequently a corm, bulb, or other variety of under- 
ground stem, as the majority of the Monocotyledons are perennial, 
herbaceous plants ; it has scattered, closed vascular bundles (Fig. 
276), and no cambium by which a continuous thickening may take 
place. The stem of the Palms, however, attains a very consider- 
able thickness, which is due to the meristem of its growing-point 
continually increasing in diameter for a lengthened period (often 
for many years), until it has reached a certain size. In this con- 
dition the growing-point has the form of an inverted cone, and it 
is only when this cone has attained its requisite size that the for- 
mation of a vertical cylindrical stem commences. Certain tree- 
like Liliaceae, as Draccena, Aloe, etc., have a continuous increase in 
thickness ; this is due to a meristematic layer, which arises in the 
cortex, outside the original vascular bundles, which were formed at 
the growing-point of the stem. This meristem continues to form 
thick-walled parenchyma and new, scattered vascular bundles. 
The primary vascular bundles, in the Palms and others, run in a 



curved line from their entrance into the stem at the base of the 
leaf, towards the centre of the stem, and then bend outwards and 
proceed downwards in a direction more parallel to the sides of 
the stem (Fig. 277). The bundles formed later, in those stems 
which increase in thickness, are not continued into the leaves. 

THE BRANCHING as a rule is very slight, the axillary buds of the 
majority of the leaves never attaining development, e.g. in the 
Palms, bulbous plants and others. As the cotyledon arises singly, 

FIG. 276. Transverse seclion of the 
stem of a Palm: v v is the wood portion, 
b b the bast portion of the vascular 

FIG. 277. Diagrammatic represen- 
tation of the course of the vascular 
bundles, from the stem into the leaves 
in a Monocotyledon. 

the succeeding leaves also must be scattered, but they are fre- 
quently arranged in two rows (Grasses, Iris, etc). The first leaf 
borne on a branch (the " Fore-leaf," 1 the bracteole, if on a floral 
shoot) has generally, in the Monocotyledons, a characteristic form 
and position, being situated on the posterior side of its own shoot, 
and hence turned towards the main axis; it is sometimes provided 
with two laterally-placed keels (Figs. 279 /, 290 0i), but the midrib 
is often absent. It arises in some cases from two primordia, which 
at the beginning are quite distinct, and thus has been regarded as 
formed by two leaves. It is, however, only one leaf, a fact which is 
evident from several circumstances, one being that it never supports 
more than one shoot, and this stands in the median plane (Fig. 279). 
THE LEAVES are amplexicanl, and have a large sheath but no 
stipules ; the blade is most frequently long, ligulate, or linear, en- 
tire, with parallel venation, the veins being straight or curved 

1 " Fore-leaf " is adopted as a translation of " Vorblatt." 



(Figs. 300, 309). Connecting the large number of veins which 
run longitudinally, there are as a rule only weak tranverse ones. 
It is very rarely that other forms of leaves are found, such as cor- 
date (Figs. 302, 312), or that the blade is branched, or the vena- 
tion is, for example, pinnate or palmate (Figs. 225, 298) ; these 
deviations are especially found in the Aracese, the Palms, the 
Scitaminefe (Fig. 308), the Dioscoreacese, and in several aquatic 
plants. The incisions in the Palm-leaf are derived by the split- 
ting of an originally entire leaf. 

THE FTRDCTURE OF THE FLOWER is generally as follows : Pr3 + 3, 
A3 + 3, G3, rarely S3 + P3 with the other members unchanged. 1 
Instead of 3, the numbers 2 and 4 may occur; rarely others. In 
all these instances there are 5 whorls, which regularly alternate 
with one another, most frequently in the 3-merous flower, as in the 
diagram (Fig. 278). This diagram is found in the following 

278. 279. 280. 

FIR. 278. Diagram of the ordinary, regular flower in the Monocotyledons : s is the 

FIG. 279. Diagram of Iris : /the bracteole ; in its axil is a shoot with its bracteole. 
FIG. 280. Diagram of Orchis : I the lip ; <r a- the two staminodes. 

orders : Liliaceas, Convallariacere, Juncacefe, Bromeliacete, Ama- 
ryllidacese, Dioscoreacese, Palmte, some Aracea?, and in some small 
orders, and may be considered as the typical structure and also 
the starting point for the exceptional orders. The ovary in many 
Monocotyledons has many ovules, and the fruit becomes a many- 
seeded berry or capsule ; this form is no doubt the oldest. In 
others the number of seeds becomes reduced to 1, and the fruit 
then becomes a cypsela, or a drupe (e.g. Graminece, Cyperacea;, 
Palmce, etc). 

Deviations from this typical floral structure in some instances 

1 Eegarding these and other abbreviations see the appendix in the book. 


may be traced to suppression, very rarely to a splitting of certain 
members, the typical relative positions not being changed. Thus, 
the Iridaceas, the Cyperacece, most of the Grramineas and some 
Juncaceas deviate in having only 3 stamens (Fig. 279), the inner 
whorl (indicated by *) not becoming developed. The Musacete 
differ in the posterior stamen not being developed ; Zingiberacere 
(Fig. 314), Marantacea?, and Cannacecv, in the fact that only 1 of 
all the stamens bears an anther, and the others are either sup- 
pressed or developed into petaloid staminodes, with some perhaps 
cleft in addition. The Orchiclea? deviate in having, generally, 
only the anterior stamen, of all the 6 developed (Fig. 280). In 
this, as in other instances, the suppression of certain parts of 
the flower is often connected with zygomorphy (i.e. symmetry in 
one plane), chiefly in the inner perianth- whorl, but also in the 
other whorls. In the Orchids, the perianth-leaf (the labellum, 
Fig. 280 /) which is directly opposite the fertile stamen, is larger 
and altogether different from the others. The perianth-leaves 
may also be suppressed; see, for example, the two diagrams of the 
Cyperaceaa (Fig. 284). In some orders the suppression of these 
leaves, which form the basis of the diagram, is so complete that it 
is hard to reduce the actual structure of the flower to the theo- 
retical type, e.g. the Grasses (Fig. 290) and Lemna (Fig. 303). In 
the first family, which especially comprises water-plants, a some- 
what different structure is found ; thus Fig. 282 differs somewhat 
from the ordinary type, and other flowers much more so ; but the 
floral diagrams which occur in this family may perhaps be con- 
sidered as the most probable representatives of an older type, from 
which the ordinary pentacyclic forms have taken their origin. In 
favour of this theory we have the larger number of whorls, the 
spiral arrangement of some of these in the flower, with a large and 
indefinite number of stamens and carpels, the perfectly apocarpous 
gynceceum which sometimes occurs, etc., etc. 

The Monocotyledons are divided into 7 Families : 

1. HELOBIFJE. This family forms a group complete in itself. It commences 
with hypogyiious, perfect flowers, whose gyucecium is apocarpous arid ter- 
minates in epigynous and more or less i educed forms. 

2. GLUJUFLORJE. These have as a starting point the same diagram as the 
following families, but otherwise develops independently. 

3. SPADICIFLOR^E. Also an independent branch, or perhaps two different 
ones which terminate in much reduced forms. 

4. ENANTIOBLAST.E. These ought perhaps to be amalgamated with the follow- 
ing family. 


5. LILITFLOR.E. These advance from forms with the typical diagram and 
bypogynous flower, to epigynous and reduced forms. 

6. SciTAiiiNE.E and 

7. GYNANDR.E. Two isolated families, which probably have taken their ori- 
gin from Liliiflorre, and have epigynous, mostly zygomorphic, and much re- 
duced forms. 

Family 1. Helobieae. 

To this family belong only water- or marsh-plants ; the endosperm 
is wanting, and they possess an embryo with a very large hypocotyl 
prolonged downwards and often club-like. The perianth is often 
differentiated into calyx and corolla ; the flower is regular, and in 
the first order-s to be considered, may be reduced to the ordinary 
Monocotyledonous type ; there are, however, usually found two 
3-merous whorls of carpels (Fig. 282), and thus in all 6 whorls, 
or again, the number of carpels maybe indefinite; the number of 
stamens also may be increased, either by the division of the 
members of a whorl, or by the development of additional whorls. 
Syncarps, 1 with nut or follicular fruitlets, are very common, for 
example, in the first orders; in the last (Hydrocharitacete) the 
carpels are not only united, but the ovary is even inferior. 

The primitive type appears to be a hypogynous flower, similar to that of the 
Juncaginacea or Alismaceas, with several 3-merous whorls, and free carpels, 
each with many ovules ; the green perianth in this instance being no doubt older 
than the coloured ones. If we take a flower with this structure as the start- 
ing point, then the family dcvelopes partly into epigynous forms, partly into 
others which are so strongly reduced and exceptional that it is scarcely possible 
to refer them to the ordinary type. The family, through the peculiar Zosteretc, 
appears to approach the Aracea?, in which Potamogetonacece and Najadacece are 
included by some authorities. However, the inclusion of Potamogeton, and with 
it Ruppia and Zannichcllui, in the Juncaginacea? appears quite correct. It would 
scarcely be right to separate Zo<t?re(e from these. Great stress has often been 
laid upon the similarity with the B-muuculacea? which is found in the Alismacea?, 
but it is scarcely more than an analogous resemblance. 

Order 1. Juncaginaceae. The $, regular, liypogijnous flowers 
have the perianth 3 + 3, sepaloid, stamens 3 + 3 (with extrorse an- 
thers), and carpels 3 + 3 (free or united), of which last, however, 
one whorl may be suppressed (in Triglochin maritima all 6 carpels 
are developed, in T.palustris the inner whorl is unfertile). Inflores- 
cence long spikes. Embryo straight. Marsh-plants with radical, 
rush-like leaves, arranged in two rows, and often sheathing and 
ligulate (" squamula3 intravaginales ") ; the inflorescence is a spike 
or raceme. Scheuchzeria. Carpels almost free ; in each at least 
1 Syncarp = cluster of fruits belonging to one flower. 



two ovules. Follicles. Triglockin has long, fine racemes without 
bracts or bracteoles ; one ovule in each carpel. The carpels in 
the two native species are united, but separate when ripe as a 
sclmocarp, loosening from below ; they open along the ventral 
suture or remain closed ; a linear central column remains. 
The most reduced is Lilcea (1-2 sp. Am.) Protogynous. About 10 species. 
Teaip. Fossils in Tertiary. 

Order 2. Potamogetonaceee. The aquatic plants belonging 
to this order are perennial, living entirely submerged, or with 
floating leaves, and preferring still water. The leaves are alter- 
nate, in some linear and grass-like, in others there is an elliptical 
floating blade, supported by a linear submerged petiole. Axillary 
scales. The fruit is generally a syncarp with nuts or drupes ; the 
embryo is curved, of very various forms. 

Potamoyeton (Pond-weed). The rhizome is creeping, sympodial 
(with two internodes in each shoot-generation) ; the inflorescence is 
a terminal, many-flowered spike, without floral-leaves ; below it are 
found 2 foliage-leaves placed nearly at the same height, from whose 
axils the branching is continued cymosely. The flowers are $ , 4- 
merous, naked, and consist only of 4 stamens, with the connectives, 
broadly developed at the back of the anthers, resembling a perianth, 
and of 4 free, sessile carpels. They are common plants in fresh water. 
The spike, during the flowering, is raised above the water. Wind-pollinated and 
protogynous. Closely allied is Pti]>pia (Tassel Pond-weed), in salt or brackish 
water. The spike has only two naked flowers, each consisting of 2 stamens and 
4 carpels. The stalks of the individual carpels are considerably prolonged. 
Zannichellia (Horned Pond-weed) is monoecious; the ? -flower consists of 4 
(2-9) carpels, with membranous, bell-shaped perianth ; long styles ; the $ -flower 
has 1 (-2) stamens. Althenia. 

Zostera (Grass-wrack) is an entirely submerged, marine plant with 
creeping rhizome (with displacement of buds) and strap-shaped 
leaves. The flowering shoots are sympodia with displacement of 
the axes (Fig. 281). The inflorescence is a peculiar, flatly-com- 
pressed spike, on one side of which the flowers are borne (Fig. 281). 
This inflorescence may be considered, no doubt correctly, to be derived from the 
symmetrical spike of Potamogeton by strongly dorsiventral development, and by 
a strong suppression of tbe floral parts taking place simultaneously. Two 
rows of flowers are developed, but of these one is so pressed into the other that 
apparently only one is present. Each flower consists of only 1 stamen 
and 1 carpel situated at the same height (Fig. 281) ; the uiiilocular 
ovary encloses 1 pendulous ovule and bears a bifid style. As 
regards the perianth (?) one leaf may be present (Z. nana, Fig. 
281 D). The pollen-grains are filamentous. Pollination takes 


FIG. 281. Zostirn. A Diagram of the branching of :he floral shoots: I, If . . . . are the 
successive shoot-geneiations, every other one being shaded ; g 1 g- .... fore-leaves; sj>> 
s])- .... spathes for the successive spikes. Each shoot is united for some distance 
with the parent axis (indicated by the half-shaded internndes-). Each shoot commences 
with a fore-leaf turning towards the parent axis, y; succeeding this is the spathe, >]> ; and 
then the inflorescence. The fore-leaf supports a new lateral shoot. B Diagram of a slioot. 
If, which is borne laterally in the axil of the fore-leaf g^, on the shoot I, g 2 its fore-leaf; 
s|i a its spathe ; sti s(iuamu]a> intravaginales. II Is the spadix with stamens and carpels ; 
b a perianth-leaf (or connective expansion, similar to those which occur in Potmnpyi'tov). 
C The upper portion of a young spadix with development of floweif . D Part of a spailix 
with 2 flowers ; the parts which theoretically belong to one another are connected by a 
dotted line. 



place under water. Ponidonia and Cynwdocea are allied to these. About 
70 species. 

OrderS. Aponogetonaceae. Aquatic plants with tuberous stem. They have 
a single, petaloid perianth (3-2-1 -leaved), most frequently 6 stamens and 3(-6) 
carpels. Straight embryo. About 15 species (Africa, Madagascar, Tropical 
Asia and Australia). Aponogeton distaclyus and A. (Ouvirandra) fenestralis are 
grown in conservatories ; the latter has lattice-like, perforated leaves. 

Order 4. Najadaceae. Only one genus Najas (about 10 species) ; annual 
fresh water plants with leaves in pairs and solitary, unisexual flowers. The $ 
flower is remarkable in having a terminal stamen, which has either 4 longi- 
tudinal loculi or 1 central OLB ; on this account the stamen of Naias is con- 
sidered by some authorities to be a stem and not a leaf-structure. The uni- 
locular gynceceum and the single, erect, anatropous ovule are also terminal. 
Pollination takes place under the water. 

Order 5. Alismaceae. The regular, hypogynous flowers are in 
some species unisexual by the suppression of either andruecmm or 
gynceceum ; they have a 6-merous perianth, generally differentiated 
into 3 sepals and 3 petals ; generally 6 stamens in the outer whorl 
(by the division of the 3 ; Fig. 282) and 
often several 3-merous whorls inside 
these, and 6- oo free carpels arranged 
cyclically or spirally. Fruit a syncarp. 
Marsh- or water-plants with radical 
leaves and long-stalked inflorescences. 

A. Butomece. Follicles with many 
seeds, irhich are borne on nearly the 
whole of the inner surface of the cyclic 
carpels (as in Nymphasacea?). Embryo 
straight. fiutomus (Flowering Rush, 

Fig. 282), has an umbel (generally composed of 3 helicoid cymes). 
(S3, P3, stamens 9 (6 + 3, i.e. the outer whorl doubled), (73 + 3. 
B. umbellatut ; creeping rhizome with triangular Iris-like leaves. Hydrocleis. 

B. Alismece. Fruit achenes. Latex common (in the inter- 
cellular spaces). The flowers are arranged most frequently in 
single or compound whorls. Embryo curved, horse-shoe shaped. - 
AUsma has S 3, P 3, A 6 (in 1 whorl, grouped in pairs, i.e. doubled 
in front of the sepals), and 1 whorl of 1-seeded achenes on a flat 
receptacle. The leaves are most frequently radicle, long-stalked ; 
the lamina have curved longitudinal veins, and a richly branched 
venation. A. plantugo. Elisma (E. natans) has epitropous (turned 
inwards) ovules, whilst the ovules of AUsma, Sagittaria and others 
are apotropous (turned outwards). Echinodorus (E. rannnculoides) 

FIQ. 282. Diagram of Bufomits .- 


has a convex receptacle, carpels many, united and capitate. 
Damasonium. Sagittaria (Arrow-head) has monoecious flowers, 
several whorls of stamens and spirally-arranged achenes on a very 
convex receptacle. S. aayittifoUa reproduces by tuberous buds formed at 
the end of long, submerged branches. The leaves, in deep and rapidly running 
water, are long and strap -shaped, but in the air arrow-shaped. 

Honey is secreted in the flower and pollination effected by insects. Alisma 
plantago has 12 nectaries. The submerged flowers of Elisma natanx remain 
closed and are self-pollinated. Biitomns has protandrous flowers. There are 
about 50 species, which mostly grow outside the Tropics. Uses insignificant. 
The rhizome of some is farinaceous. 

Order 6. Hydrocharitaceae. This order differs chiefly from 
the preceding in its epigynous flowers. These are in general uni- 
sexual {dioecious}, and surrounded by a 2-leaved or bi-partite 
spat he ; they are 3-merous in all whorls, but the number of whorls 
is generally greater than 5, sometimes even indefinite. The peri- 
anth is divided into calyx and corolla. The ovary is unilocular with 
parietal placentation, or more or less incompletely plurilocular. The 
fruit is berry-like, but usually ruptures irregularly when ripe. Em- 
bryo straight. Most often submerged water-plants, leaves seldom 
floating on the sm-face. Axillary scales (squamulce infravaginales). 
Hydrocharis. Floating water-plants with round cordate leaves ; 
S3, P3 (folded in the bud); ^-flowers : 3 (-more) flowers inside 
each spathe ; stamens 9-15, the most internal sterile. $ -dowers 
solitary ; three staminodes ; ovary 6-locular, with many ovules 
attached to the septa ; styles 6, short, bifid. [The petals of the 

9 -flowers bear nectaries at the base. In this and the following 
genus the pollination is without doubt effected by insects.] 
H. morsus ranee (Frog-bit) has runners; it hibernates by means of special 
winter-buds. Stratiotes; floating plants with a rosette of linear, 
thick, stiff leaves with spiny margin, springing from a short 
stem, from which numerous roots descend into the mud. In- 
florescence, perianth, and ovary nearly the same as in Hydrocharis, 
but the (J -flower has 12 stamens in 3 whorls, of which the 
outer 6 are in 1 whorl (dedoublement), and inside the perianth 
in both flowers there are numerous (15-30) nectaries (stami- 
iiodesP). (S. aloides (Water-soldier); in N. Eur. only $ -plants. 

Vallisneria i-ph-uli* is a tropical or sub-tropical plant, growing gregariously on 
the mud in fresh water. The leaves are grass-like, and the plants dioecious ; 
the $ -Mowers are detached from the plant, and rise to the surface of the water, 
where they pollinate the <j> -flowers. These are borne on long, spirally-twisted 
peduncles which contract after pollination, so that the ? -flower is again drawn 
under the water, and the fruits ripen deeply submerged. Elodea canadensis 


is also an entirely submerged plant. The leaves are arranged in whorls on a 
well-developed stem. Only $ -plants in Europe (introduced about 183G from 

N. Am ). This plant spreads with great rapidity throughout the country, the 
reproduction being entirely vegetative. Hydrilla, Haliphila, Thalassiu,Enhalus. 

In many of these genera the number of whorls in the flower is remarkably re- 
duced; for example, in Vallisneria, in the <? -flowers to 2: Pr 3, A (1-) 3, in the 

? to 3 : Pr 3, Staminodes 3, G 3. About 40 species ; Temp, and Trop. 

Family 2. Glumiflorae. 

The hypogynous flowers in trie Juncaceas are completely developed 
on the petitacyclic, trimerous type, with dry, scarious perianth. Even 
in these the interior whorl of stamens becomes suppressed, and the 
ovary, which in Jnncus is trilocular with many ovules, becomes in 
Lnzula almost uniloculai-, but still with 3 ovules. The perianth in 
the Cyperaceae and Gramineae is reduced from hairs, in the first of 
these, to nothing, the flowers at the same time collecting- more 
closely on the inflorescence (spike) supported by dry bracts (chaff) ; 
the number of stamens is almost constantly 3 ; stigmas linear; the 
ovary has only 1 loculus with 1 ovule, and the fruit, which is a cap- 
sule in the Juncacese, becomes a nut or caryopsis. The endosperm 
is large and floury, the embryo being placed at its lower extremity 
(Pigs. 286 B, 291). The plants belonging to this order, with the 
exception of a few tropical species, are annual or perennial herbs. 
The stems abore ground are thin, and for the most part have long 
internodes, with linear, parallel-veined leaves which have long 
sheaths, and often a ligule, i.e. a membranous projection, arising 
transversely from the leaf at the junction of the sheath and blade. 
The underground stems are short or creeping rhizomes. The 
flowers are small and insignificant. Wind- or self-pollination. 

Order 1. Juncaceae (Rushes). The regular, hermaphrodite, 
hypogynous flowers have 3 + 3 brown, 
dry, free perianth - leaves projecting 
like a star during the opening of the 
flower ; stamens 3 + 3 (seldom 3 + 0) 
and 3 carpels united into one gynceceum 
(Fig. 283) ; the ovary is 3- or 1-locular ; 
there is as a rule 1 style, which be- 
comes divided at the summit into 3 
stigmas, often bearing branches twisted 
to the right (Fig. 283). Fruit a capsule 

with loculicidal dehiscence. The embryo is an extremely small, 
ellipsoidal, cellular mass, without differentiation into the external 


Jnncus (Rush) has glabrous foliage-leaves, generally cylindrical, 
rarely flat ; the edges of the leaf-sheath are free (" open " leaf- 
sheaths) and cover one another. The capsule, 1- or 3-lbcular, 
with many seeds. Luzula (Wood-Rush) has flat, grass-like leaves 
with ciliated edges ; the edges of the leaf-sheath are united 
("closed " leaf-sheath). The capsule unilocular and '3-seeded. 
Prioninm : S. Africa ; resembling a Tacona. 

The interior whorl of stamens, in some species, disappears partially or 
entirely (J. supinus, capitatus, conglomeratus, etc.) 

Some of the numerous Juncus-species (e.g. J. eff'usus, glauctiK, conglomeratw, 
etc.), have false, lateral inflorescences, the axis of the inflorescence being pushed 
to one side by its subtending leaf, which apparently forms a direct continua- 
tion of the stem, and resembles it both in external and internal structure. 
The foliage-leaves of this genus were formerly described as " unfertile stems,'' 
because they are cylindrical, erect, and resemble stems, and consequently the 
stem was said to be " leafless " : J. eff'it-su-s, ylaucus, conglameratus. Stellate 
parenchymatous cells are found in the pith of these sterns and in the leaves. 
Other species have distinct terminal inflorescences and grooved leaves ; J. 
b 11 fun ius (Toad-rush), compressus, and others. The inflorescences most often 
present the peculiarity of having the lateral axes protruding above the main 
axis. Their composition is as follows : The flowers have either no bracteoles, 
and the inflorescences are then capitulate ; or they have 1-several bracteoles. 
Each branch has then, first, a 'J-keeled fore-leaf placed posteriorly (" basal- 
leaf "), and succeeding this are generally several leaves borne alternately and in 
the same plane as the basal-leaf, the two uppermost (the " spathe-lea^es '') being 
always barren ; those which lie between the basal-leaves and the spathe-leaves 
are termed ' intermediate-leaves." If only branches occur in the axils of the 
basal-leaves, then the succeeding branches are always borne on the posterior 
side of the axis, and form a fan * ; if the basal-leaf is barren, and if there 
is only one fertile intermediate-leaf, then the lateral axes are always on tbe 
upper side, and a sickle'-like inflorescence occurs ; if there are 2 fertile 
intermediate-leaves, then a dichasium is formed, and in the case of there 
being several, then a raceme, or spike. 

Jtmcacea; are, by several authors, classed among the Liliiflorse, but there are 
so many morphological and partly anatomical features agreeing with the two 
following orders, that they may, no doubt, most properly be regarded as the 
starting point of these, especially of the Cypemcece, which they resemble in the 
type of flowers, the inflorescence, the type of mechanical system, and the stomata. 

POLLINATION by means of tlie wind. Cross-pollination is often established by 
protrgyny. J. bufonius has partly triandrous and cleistrgamic, partly hexan- 
drous, open flowers. DISTRIBUTION. The 200 species are spread over the entire 
globe, but especially in cold and temperate countries ; they are seldom found in 
the Tropics. USES. Very slight ; plaiting, for instance. 

Order 2. Cyperaceae. The majority are perennial (seldom 

" Fan " and " sickle " are adopted as terms for these inflorescences from the 
German "fiichel" and '' sicliel." 




FIG. 28i. DiRgram of structure of: 
A Soii-pus siluaticus ; -B Eriophorum 

annual) herbs living in damp situations, with a sympodial rhi/ome 
and grass-like appearance. The stems are seldom hollow, or have 
swollen nodes, but generally triangular, with the tipper internode 
just below the inflorescence generally very long. The leaves are 
often arranged in 3 rows, the leaf-sheath is closed (very seldom 
split), and the ligule is absent or insignificant. The flowers 
are arranged in spikes (spikelets) which may be united into other 
forms of inflorescences (chiefly spikes or racemes). The flowers 
are supported by a bract, but have 
no bracteoles. In some genera the 
perianth is distinctly represented 
by six bristles corresponding to 
six leaves (Figs. 284 A, 286 A} ; 
in others it is represented by an 
indefinite number of hairs (Fig. 
284 B), and very frequently it is 
altogether wanting. The inner 
u-horl of stamens is absent, and the flower has therefore 3 stamens 
(rarely more or less than 3), the anthers are attached by their bases 
to the jilnment (innate) and are not bifid (Figs. 286). Gynoeceum 
simple, formed of 3 or 2 carpels ; 1 style, which is divided at the 
extremity, as in the Juncacea?, into 3 or 2 arms ; the single loculus 
of ihe ovary contains one basal, erect, anatropous ovule; the 
stigmas are not feather-like. Fruit a nut, whose seed is generally 
not united with the pericarp. The embryo is small, and lies at 
the base of the seed in the central line, surrounded on the inner side 
by the endosperm (Fig. 286 7>). On germination the cotyledon 
does not remain in the seed. 

A regular perianth, with 6 scale-like perianth-leaves in 2 whirls, is found in 
Oreoboluf. In Scirptis littoralis the perianth-leaves are spreading at the apex, 
and divided pinuately. 

The branching of the inflorescence is o f ten the same as in the Juncaceae, and 
supports the theory that these two orders are related. In Rhynchospora and 
others, the "spikelets" are really only "spike-like" and to some extent compound. 


1. Spikelets cylindrical, the bracts arranged spirally (in 
many rows). The lower ones are often barren, each of the others 
supports a flower. Scirpus (Club-rush). The spikelets are 
many-flowered; the perianth is bristle-like or absent, and does not 
continue to grow during the ripening of the fruit (Fig. 286 A). 
Closely allied to this is Heleocharis, with terminal spikes. 



Eriophornm (Cotton-grass) differs chiefly in having the perianth- 
hairs prolonged, and forming a bunch of white, woolly hairs (Fig. 
284 B). 

Clndium and Rlnjnchospora (Beak-rush) differs especially in the/eze-flowered, 
compound spikelets which are collected into small bunches ; the latter Las 
received its name from the fact that the lowermost portion of the style remains 
attached to the fruit as a beak. 

2. Spikelets compressed, the bracts arranged only in two rows ; 
the other characters as in the first-mentioned. Cyperus (spikelets 
many-flowered) ; Schcenus (Bog-rush) ; spikelets few-flowered ; 
S. nigricans has an open sheath. 


In the ^-flowers there is no trace of a carpel, and in the ? 
no trace of a stamen. Floral-leaves in many rows. In some 
(Scleria, certain Careoi-species) , g - and ? -flowers are borne in the 
same spikelet, the latter at the base or the reverse ; in the majority 
each spikelet is unisexual. 

Gar ex (Fig. 285) has naked, most frequently monoecious flowers. 
The $ -spikes, which are generally placed at the summit of the 
whole compound inflorescence, are not compound ; in the axil of 
each floral- leaf (bract) a ftoiver is borne, consisting only of a short 

285. 2S6. 

FIG. 285. Cares : A diagram of a male flower ; B of a female flower with 3 stigmas ; C of 
a female flower with 2 stigmas; D diagrammatic figure of a female flower; E similar one of 
the androgynous (false) spikelet of Elyna. The <J is here represented placed laterally; it 
is terminal, according to Pax. 

FIG. 286. A Flower of Scirpus lacustna. B Seed of Carex in longitudinal section. 

axis with three stamens (Fig. 285 A). The ? -spikes are compound ; 
in the axil of each floral-leaf is borne a very small branch (Fig. 
285 D, a) which bears only one leaf, namely, a 2-keeled fore-leaf 


(utriculus, ntr. in the figures) which is turned posteriorly (as 
the fore-leaves of the other Monocotyledons), and being obliquely 
sheath-like, envelopes the branch (in the same manner as the sheath 
of the vegetative leaves), and forms a pitcher-like body. In the 
axil of this leaf the ? -flower is situated as a branch of the 3rd 
order, bearing only the 2-3 carpels, which are united into one 
gynoeceum. The style protrudes through the mouth of the utri- 
culus. The axis of the 2nd order (a in Fig. 285 D) may sometimes elongate as 
a bristle-like projection (normally in Uncinia, in which it ends as a hook, 
hence the uame) ; this projection is in most cases barren, but it sometimes 
bears 1-several bracts which support male-flowers ; this is normal in Elyna 
(or Kobresiu) and Schasnoxiphium ; the axis ( in '285 E) bears at its base a 
female-flower supported by the utriculus, and above it a male-flower supported 
by its bract. 

POLLINATION by means of the wind. Protogynous. Sometimes self-pollinated. 
The order embraces nearly 3,000 species, found all over the world. Care.c 
and Scirpiis are most numerous in cold and temperate climates, and become 
less numerous towards the equator. The reverse is the case with Cy perns 
and other tropical genera. They generally confine themselves to sour, swampy 
districts; some, on the other hand, are characteristic of sand-dunes, such as 
Sand-star (Car ex arenaria). There are about 70 native species of Carex. 

USES. In spite of their large number, the Cyperacese are of no importance 
as fodder-grasses, as they are dry and contain a large amount of silica ; hence 
the edges of many of the triangular stems or leaves are exceedingly sharp 
and cutting. Cyperus escitlentiis has tuberous rhizomes, which contain a large 
amount of fatty oil and are edible (earth-almonds) ; it has its home ia the 
countries of the Mediterranean, where it is cultivated. 

Cyperus ptipynis (W. Asia, Egypt, Sicily) attains a height of several metres, 
and has stems of the thickness of an arm which were used by the ancient 
Egyptians for making paper (papyrus). Some serve for plaiting, mats, etc. 
(Scirpus laeustris, etc.). Isolepis is an ornamental plant. 

Order 3. Gramineae (Grasses). The stems are cylindrical, 
generally hollow with swollen nodes, that is, a swelling is found at 
the base of each leaf which apparently belongs to the stem, but in 
reality it is the swollen base of the leaf. The leaves are exactly 
alternate ; the sheath is split (excep. Broimis-species, Poa pratensis, 
P. trh-ialis, Melica, Dactylis, etc., in which the sheath is not split), 
and the edges overlap alternately, the right over the left, and vice 
versa; the ligule is nearly always well developed. In general, the 
flowers are hermaphrodite ; they are borae in spikelets with alternate 
-floral-leaves, and the spikelets themselves are borne in either 
spikes or panicles. The two (seldom more) lowest floral-leaves in 
each spikelet (Fig. 289 0Y, nY) are barren (as the covering-leaves 
in many umbels and capitula) ; these are termed the glumes. The 



succeeding floral-leaves, each of which supports one flower as its 
bract, are called the outer pales (nl) ; these sometimes each bear an 
" awn " (a bristle-like body which projects in the median line either 
from the apex or the back) ; sometimes the upper ones are barren. 
Each flower has a brac.teole, which is placed on the inside opposite 
the main axis ; it is thin, binerved or two-keeled, and never has an 
awn; it is known as the inner pale (01). Immediately succeeding 
the bracteole are : (<?) some small, delicate scales (lodicules, Figs. 
287 D, 288 C, 290 L) ; (6) three stamens with anthers versatile, 
so as to be easily moved, and usually notched at each end (Fig. 

287 (7) ; and (c) a simple gyncecenm formed of one carpel with tivo 
styles having generally spirally -branched stigmas (Figs. 287 D, 

288 C). The ovary is unilocular, and contains one ascending or 
pendulous, anatropous ovule. Fruit a nut, whose seed is always 
firmly united with the thin pericarp ("caryopsis "). The embryo is 

FIG. 287. Triticum : A axis (rachis) of ear showing the notches where the spikelets 
were inserted; Ban entire spikelet; Ca flower with the pales; Da flower without the 
pales, showing the lodicules at the base ; E glume ; F outer pale ; G inner pale ; H fruit; 
/ longitudinal section of fruit. 

larger than in the Cyperacere and is placed at the base of the seed, 
but on the otiter convex surface of the pericarp (Figs. 287 I, 288 



288 -D, 291), outside the endosperm ; plumule large with several 
leaf-primordia. On germination the cotyledon remains in the seed. 
The majority of Grasses are annual or perennial herbs ; tree- 
like forms being only found in the Tropics, for example, the 
Bamboos; they branch (in tufts), especially from the axils of the 
basal-leaves, while those which are borne higher on the stem are 
separated by longer internodes and have no vegetative branches in 

Fto. 289. Diagramatic outline of 
a spikelet : n Y lower glume ; o Y 
upper glume ; n I upper pale ; o I 
the inner pale; I-Z lodicules ; s-t 
stamens; I-I main axes; II lateral 

FIG. 283. B co in us moIUs : A inflorescence ; 
B the uppermost flower of a spikelet, with 
its axis turned forward ; in front is seen the 
two-keeled inner pale (bracteole) and the 
stamens protrude between this and the outer 
pale (.bract) ; C a.n ovary with the 2 stigmas 
oil its anterior side, the 2 lodicules, and the 
3 stamens ; D the fruit seen from the dorsal 
side; E the same from the ventral side. 

their axils, though a few forms, like Bambusa and Calamagrostis 
lanceolate^ produce branches in these axils. 

Only a few Grasses have a solid stem, such as Maize, Sugar-cane, and Andro- 
pogon. The blade is flat in the meadow-grasses, but the Grasses which live on 
dry places (" prairie-grass ") exposed to the sun, often have the blade tightly 
W. B. U 



rolled up and almost filiform or bristle-like, with anomalous anatomical 
structure. A closed tubular sheath is found in Helica uni flora, B;-onn(s-species, 
Poa pratensiis and trivialis, Briza and some Gif/cmYt-species. The sheath is 
developed for the purpose of supporting the young internodes while their growth 
is proceeding at the base. The "nodes" (the swollen joints which are seen on 
stems of Grasses) are not really part of the stem but are formed by the base of 
the leaf-sheath. They play a part in assisting the haulms to regain a vertical 
position when laid prostrate by wind or rain. The awn on the pale is homo- 
logous with the blade of the Grass-leaf, and the pale itself is the sheath. The 
arrangement of the leaves in the spikelet is similar to that in Cyperus and other 
Cyperacea?, their floral-leaves being borne in several rows in Streptochceta. More 
tbau two barren "glumes " are found in Streptochata, several Phalaridea? and 
others. The spikelets, too, are again arranged in two rows in the axils of sup- 
pressed floral-leaves. The inflorescence becomes a " compound spike " (ear) when 

FIG. 290. Diagram of the Grass-flower : 
ni outer pale ; en inner pale ; l-l lodi- 

FIG. 291. Longitudinal section of an 
Oat-grain : a the skin (pericarp and 
testa); b the endosperm; c the cotyledon; 
d the plumule. 

the spikelets are sessile. In the majority of instances the spikelets are borne 
on long stalks ; when these branch, then the secondary branches, and similarly 
all branches of higher order, are placed so far down upon the mother-axis that 
they all appear to be of equal value and to arise in a semicircle from the mother- 
axis itself, though in reality they arise from each other (Panicle, Fig. 288 A). 
Sometimes the main axis and branches of different orders unite together as in 
Al ipecnrus. Phleum, and some other Grasses, and hence the single (short- 
stalked) spikelets appear to arise singly and spirally, or without any definite 
order, directly from the main axis, with the production of a cylindrical inflor- 
escence bearing " spikes " OH all si,lex, that is, a " strike-like panicle. " Many 
inflorescences are somewhat dorsiventral. The flower is rarely unisexual (Zea 
mats) or barren. Considerable difficulty is experienced in reducing the Grass- 


flower to the ordinary 3-merous Monocotyledonous type. Some authorities 
consider the lodicules, which are present in all Grasses but absent in the 
Cyperaceas, to be homologous with a perianth. According to a more recent 
theory they are bracteoles, and hence the Grarninefe, like many oi the Jun- 
caceffi, have 2-3 bracteoles placed in two rows in the median plane. li this 
theory be correct, the floiver is naked. The lodicules expand quickly and 
cause the opening of the flower (i.e. the two pales become separated from each 
other). Generally only 3 stamens belonging to the outer whorl are present (Fig. 
290), as in Ii is (Fig. 279), certain Juncacere and Cyperacete (Fig. 281), but in 
some, such as the Rice and certain species of Bamboos, all 6 are found. 
Pariaiia has more than 6. Only 1 of the carpels is present, namely, the 
anterior (of those in Fig. 281), so that the ventral suture and the place of at- 
tachment of the ovule are situated at the back of the ovary. The number of 
styles does not correspond with the number of carpels, and the styles may 
therefore be supposed to arise from the edges of the leaf to the right and left a 
position which is not without analogy. In addition, a stylar projection is some- 
times found on the anteiior side and in the median line (e.g. in Phragmites), 
and the solitary style iu Nardus has exactly this position ; a similar arrange- 
ment is found in some species of Bambino, which have only one style; other 
species of liambiisa have three styles. A tripartite style is found in Phanis. 

[The Grass-flower may be reduced to the ordinary Monocotyledonous type 
thus: The outer pale is the bract of the flower since it bears iu its axil the 
floral shoot ; the inner pale occupies the customary position of the bracteole. 
The fact that it is binerved can be explained by its having been pressed against 
the main-axis during development. Similar biuerved bracteoles are found in 
7m (Fig. 279). These bracteoles in both Grass and Iris arise from single pri- 
mordia, and are not produced by the coalescence of two leaves. The lodicules 
are the only parts of the perianth remaining, the outer whorl having been sup- 
pressed, and also the posterior leaf of the inner whorl ; a posterior lodicule, 
however, is found in the Rice and some species of Bamboo. The outer whorl of 
stamens is usually absent, though this again is present in the Rice and Bamboo. 
The three carpels are reduced to one with two or sometimes three stigmas.J 

THE FLOWERING. In the panicles the flowers open in basipetal order; 
the flowers in the spikes situated somewhat above the middle, commence to 
open first, and the flowering proceeds upwards and downwards. A few Grass- 
flowers never open (cleistogamic) ; Leersia oryzoid's, Stipa-species, and e.g. 
Wheat and Rye in cold damp weather ; some open their pales so wide that the 
anthers and stigmas may protrude at the top ; most frequently the lodicules 
expand and force the pales suddenly and widely apart. The filaments elongate 
considerably, so that the anthers are pendulous and the stigmas unfold. la 
some Grasses Wheat, the blooming of each flower only lasts a short time. 
POLLINATION is generally effected by the wind. The Eye separates the pales 
very widely in the morning, and allows the anthers and stiguaas to appear ; 
it is almost entirely sterile when self-pollinated. The Wheat flowers at any 
time of the day, each flower lasting only a quarter of an hour. The pales open 
suddenly, but only half way, and the anthers scatter one-third of the pollen in 
their own flower and two-thirds outside. Self-pollination is effectual, but 
crossing gives better results. In Hordeum vulgare (all flowers $ ) the flowers 



of the 4 outer rows behave as in the Wheat, but those in the two central rows 
always remain closed. The -flowers in the two central rows of H. distichtim 
remain closed and fertilise themselves ; they open exceptionally, and may be 
pollinated by the c? -flowers in the 4 lateral rows. H. liexastichum is cleisto- 
gamic. Ofitis pollinate themselves. 

The ripe Grass-fruit, in some species of Bamboo, is a berry; in some other 
Grasses a nut with loosely lying seed, in some even a capsule, but otherwise a 
"caryopsis." In some instances it is loosely enveloped by the pales (Oat), in 
others firmly attached to these (Barley), and finally, in others, "naked," i.e. it 
is entirely free from the pales (Wheat and Rye). On the ventral side there is a 

FIG, 292. Barley grain : A sec- 
tion through the skin (a-d) and the 
most external part of the endo- 
sperm ; 61 the " aleurore layer" ; 
st starch-containing cells; B starch 

FIG. 293. Wheat- 
grain germinating: g 
the plumule; b the first 
leaf succeeding the co- 
tyledon ; r 1 the primary 
root ; r~ lateral root. 

FIG. 291. -Older seed- 
ling of the Wheat: sthe 
second sheathing-leaf ; 
I first foliage-leaf. 

groove (Fig. 288 E) ; on the anterior side (dorsal suture), which is turned towards 
the inner pale, it is convex, and at the base on this side, inside the testa, lies the 
embryo (Fig. 288 D). The apex of the fruit is often hairy (Fig. 293). The xA-m 
(Fig. 291, a) is formed by the pericarp and testa, and in some cases (Barley) the 
pales also form the outer portion. The endosperm (b) is large, and formed of 
pareiichymatous, starch-containing cells : aleurone (proteid) grains may also be 
found among them. When the starch-grains and the aleurone-grains adhere 
together the endosperm becomes " horny," but is " floury " when the starch- 
grains lie loosely with air between them. In the most external region, just 
beneath the skin, 1-several layers of nearly cubical cells (filled principally with 
fileurone-grains and fat) are found, the aleitrone-layei' (Fig. 292). The embryo 
(Fig. 291 c-d) contains large quantities of fatty oil ; the large shield-like structure, 
attached to the embryo and turned inwards towards the endosperm (c), is the 
cotyledon (scutellum) ; it remains enclosed in the seed during germination, and 


dissolves the endosperm by means of the peculiar epithelial cells developed on 
the dorsal surface. The radicle, on germination, is obliged to perforate a mass 
of cells derived from the suspensor and which form the "root-sheath" 
(coleorhiza, Fig. 293) round its base. In addition to the tap-root, lateral roots 
are frequently developed before germination ; these quickly break through, 
and later on are followed by others which appear at the base of the leaf (Figs. 
293, 294). 

The DISTRIBUTION OF THE FRUIT is most frequently effected by the wind. The 
spirally-twisted and hygroscopic awn which persists on the fruits of some species 
(Arena, Stipa, etc.) assists in their dissemination, and eveia helps to bury them 
in the ground. 

The two preceding orders are more closely related to each other than they 
are 10 the Grammes. 

The generic differences are chiefly founded 011 the form of the inflorescence, 
the number and sex of the flowers in the spikelets, the shape and relative 
length of the pales, awns, etc. In addition to these the structure of the fruit 
audsted presents a great many differences ; some have compound starch-grains, 
while in others they are single ; some have 1 layer of aleurone-cells, others have 
several (Fig. 292), etc. 

1. BAMBUSE.E. Tall Grasses with woody, very siliceous stems 
which bear many brandies in the axils of the leaves. 6 stamens. 
Bambusa (Bamboo). 

2. OiiYZEjE. Onjza saliva (Rice) is a herbaceous marsh-plant, 
with panicle and small, 1-flowered spikelets, with two small glumes 
and two large, boat-shaped, strongly siliceous pales. 6 stamens. 

Leersia. Lygeuni. Pharus. Zizania aquatica. 

3. MAYDE^E. Zea mais (Indian-corn, Maize) ; the spikelets are 
unisexual ; the <J -spikelets in a terminal panicle ; the ? -spikelets 
closely crowded and arranged in many rows in a thick, axillary 
spike, enclosed by large sheathing-leaves. The ? -spikelets are 
l-(2-) flowered ; the ovary bears one, long, filamentous style, with 
bind stigma. Euchl&na; Coix. 

4. ANDROPOGONEJ;. Saccharum (Sugar-cane) ; the spikelets are 
exceptionally small, 1-flovvered, and borne in pairs in many- 
flowered, long-haired panicles. Tall grasses with solid, sappy 
stem. Andropoyon. 

5. FESIUCEJC. Grasses with panicle (or spike-like panicle) and 
2-several-flowered spikelets. Glumes small, in each case shorter 
than the spikelet. Festaca (Fescue) and Bromas (Brome, Fig. 
288) have the awn placed at the apex of the pale, or slightly 
below it. Fettnca has perennial species, with only a sparsely- 
branched panicle with branches solitary or in pairs, and round 
spikelets ; the leaf-sheath is widely open. Bromus has the 


branches borne in half whorls, and the leaf-sheath scarcely half 
open. Brachypoilinm has very short-stalked spikelets in a raceme. 
Poa (Meadow-grass), Briza (Quaking-grass) and Glyceria have 
awnless spikelets ; these in Poa are ovoid, compressed, and with 
sharply-keeled glumes; in Briza they are broad, cordate and 
drooping, with boat-shaped glumes ; in Glyceria round, long, many- 
flowered, linear or lanceolate : some species of Glyceria have closed 
leaf-sheaths.' Dactylis (Cock's-foot) differs from all others in the 
somewhat crowded and unilateral (subsecund) spikelets, which are 
compressed and oblique (i.e. one side more convex than the other). 
Phragmites (P. communis, Reed); the lowermost flowers of the 
spikelet are <$ ; its axis is covered with long, silky hairs ; pales 
without awns, but acuminate. Perennial marsh-plants. Melica ; 
panicle small, sparsely-branched with round, awnless, few-flowered, 
usually drooping spikelets. The upper pales, with arrested flowers, 
are generally united into a club-like mass. Mulinia, Eragrostis, 
Koeleria, Catabrosa. Cynosurus (Dog's-tail) has a small, spicate 
panicle with unilateral spikelets, some of which are fertile, some 
barren, each supported by a pectinate scale. Arundo. Sesleria. 
Gi/nerium. Triodia. 

6. AVENE.E. Panicles with 2-many-flowered spikelets ; at least 
one of the glumes is quite as long as the entire spikelet. 
Arena (Oat). The pale is boat-shaped, often bifid, and at about 
the middle of the back has a twisted, bent awn. Air a (Hair-grass) 
has a long-branched panicle with small, 2-flowered spikelets ; the 
pale has a dentate apex and bears an awn on the posterior side 
close to the base. Weingilrineria. Holcus (Yorkshire-fog) ; a soft, 
hairy Grass with an open panicle, keeled glumes ; 2 flowers in the 
spikelet, of which the lower one is $ , the upper <$ ; the pale 
which supports the ty -flower has no awn, but that which supports 
the (J -flower, on the contrary, is awned. 

7. AoROSTlDEyE. Panicles or spike-like panicles with 1 -flowered 
spikelets. Generally 2 glumes and only 1 pale. The following 
have PANICLES : M ilium with square panicle-branches and round 
spikelets; Agrostis (Fiorin), with compressed, glabrous spikelets, 
whose glumes are longer than the pales. Calamagrostis differs in 
having a chaplet of long hairs at the base of the pale. Sti^a 
(Feather-grass) has a long, twisted awn. The following have SPIKE- 
LIKR PANICLES : Plileum (Cat's-tail, Timothy-grass) has sharply 
pointed, entirely free glumes, which are much longer than the 
awnless pales. Alopecnrus (Fox-tail) ; glumes united below ; pale 


with awn. Ammophila (Psammct). A. arundiuacea; pales hairy 
at base; perennial, stiff- leaved, glaucous sand-grass with creeping 
rhizome. Aristida. Sporobolus. 

8. PHALARIDE^:. Panicles and spike-like panicles. The spike- 
let has in the upper part a single fertile flower ; below it are 
placed 4 pales, of which the upper 1-2 sometimes support - 
flowers. On the whole, 6 floral-leaves of the first order are 
present. Phalaris (P. canariensis, Canary-grass) has an ovate, 
spike-like panicle, the spikelets are compressed, convex on the 
outer side, concave on the inner. The large glumes are winged 
on the back. Digraphis (D. anindinacea) is closely allied to 
Phalaris, but the keel of the glumes is not winged. AnthoxantTium 
(A. odoratum, Sweet-vernal) has a small, lanceolate, open, spike-like 
panicle; the spikelets have below 2 barren flowers, and a.bove 
these an -flower with 2 stamens. The upper glume is longer 
than the flower. Hierochloa. 

g. CHLOEIDE.E. The spikelets are arranged in the form of a spike in two 
rows on one side of an often flatly-compressed axis ; they are mostly 1-flowered. 
Cliloris ; Cteiiinm ; Cynodon ; Eleusine ; Jlicrochloa. 

10. PANICE.E. The spikelets are borne in panicles or spikes, which may be 
arranged like fingers or in a raceme. There is a centrally-placed $ -flower ; 
below it is sometimes a $ -flower. Panicum ; Paapalum ; 02)lismenus ; Setaria 
has an almost cylindrical spike-like panicle with several barren brauchlets, 
which project as stiff, rough bristles. Cenchrus ; Pennisetum. 

II. HORDED. Spikes compound ; spikelets sessile in the notches 
of a toothed axis. 

A. Spikelets solitary. Triticum (Wheat, Fig. 287) has in each 
tooth of the main axis, a several-flowered spikelet which turns its 
flat side towards the central axis. The cultivated species (tine 
Wheat) are 1-2-annual, the wild ones (T. repens, Couch, also as 
an independent genus, Agropyrum) are perennial, with creeping- 
rhizome and lanceolate glumes. Lolium (Rye-grass) has in each 
tooth of the main axis a many-flowered, compressed spikelet, 
which is placed edgeivise towards it and (with the exception of 
L. perenne) has only one outwardly-turned glume (L. temulentum 
has a rudiment of the inwardly-turned lower glume) ; the terminal 
spikelet has two glumes.' Secale (Rye). A two-flowered spikelet 
in each tooth ; small, lanceolate, acuminate glumes. Nardus and 
Lepturus have very narrow spikes, the former with unilateral 

B. In each notch of the axis 2 or more spikelets are placed 


close together. Hordenm (Barley). In each tooth three 1-flowerecl 
spikelets. H. hexast it-hum (6-rowed Barley), has 6 rows of fruits, 
since all the spikelets'are fertile, and H. distichum (2-rowed Barley) 
2 rows, since the lateral spikelets are <$ , and barren (p. 292). 
Elymus has 2-6 many-flowered spikelets in each joint of the main 
axis. JEgilops has awns upon the glumes. 

DiSTKiBtiTiON. 315 genera with 3,500 species. The order is distributed 
over the whole world, and as regards number of individuals is perhaps the 
richest. In the Tropics, large, broad-leaved, tree-like forms are found (Bam- 
l>use<e, Olyreff, Andropogonete, etc. ; in S. Europe, Amiido doita.r) ; in England, 
next to the Composite?, it is the order most rich in species (about 134). The 
origin of some of the cultivated Grasses is lost in obscurity. The Maize, no 
doubt, was indigenous to America, where its nearest relatives are found, and 
where it has also been discovered in ancient Indian graves ; Durra or Guinea- 
corn, Millet and Sugar-cane are South Asiatic plants, and our own cereals no 
doubt have sprung primarily from Western Asia and South-Eastern Europe 
(Barley from Armenia and Persia, where a very closely related wild species 
is found ; Wheat from the same districts ; Rye from the perennial species 
.S. wontanum). Panicum altissimum and Eice have come from Africa. 

USES. The Grasses play a very important part as cereals and fodder plants. 
The following are the most important of the cultivated ones: Triticum rulgare 
(common Wheat), turgidum, amultntm, polonicum, fpelta, dunini, etc. ; Secalc 
cerr.ale (Rye) ; Barley (Horcleiun-species, see under the genus) ; Maize ; Oats 
(Avena saliva, oricntalis, inida) ; Millet (Panicum miliaceum) ; Durra (Turkish 
Millet, or Guinea corn, Sorghum vulyare, cernuum and saccharatuni) ; Manna- 
grass (Glyceria fliuton*). As fodder-plants especially: Rye-grass (Lolium 
perenne); Oat-grass ( elatior) ; Timothy (Plileum pratem?) ; Fox-tail 
(Alopecurus pratemis) ; Cock's foot (Dactylic glomrrata) ; Dog's tail (Ci/nominat 
cristatus); Sweet-vernal (Antlioxanthum odoratum) ; Soft grass, or Yorkshire-fog 
(Holcus lanatus and mollis) ; Quaking-grass (Briza mdia) ; species of Meadow- 
grass (Poa) ; Fescue (Fcstnca) and Brome (Bromiii). Several cultivated species 
of Grass are also used in the preparation of fermented liquors, the starch in 
the seeds being transformed to sugar (beer from " Malt," i.e. the germinated 
Barley ; arrack from Rice) ; or the stem becomes specially saccharine before 
flowering: the Sugar-cane, Sorghum saccharatuni. 

OFFICINAL. The rhizome of Triticum repem, Oat-grain, flour of Barley, and 
the starch of Wheat, also sugar. 

The seeds of Lolium temulentum are considered 2^oisonous. The stems of 
many species (including our common grains) are used in the manufacture 
of paper, especially "Esparto grass" (Sti2>a tenacissima) from Spain and N. 
Africa, and the sheathing-leaves of the ? -spike of Maize. Sand Lyme-grass (Ely- 
mus arenarins), and especially Psamma arenaria, are important. But few Grass- 
species are sweet-scented : Anthoxanthum odoratum and Hierochloa cdorata 
contain cournarin ; Andropogon-species have essential oils (' Citronella oil"). 
ORNAJIENTAL PLANTS are : the " Ribbon-grass" (a variety of Diijruphis arundi- 
nacca), Stipa peunata (whose awn is exceedingly long and feathery), Gynerium 
argenteum (Pampas-grass), Lagurus oratus, Hordeum jubatitm, Lronuts Irizi- 



Family 3. Spadiciflorae. 

The primitive form resembles that of the preceding family. In 
it we find the typical, perfectly developed, Monocotyledonous 
flower, sometimes even with free carpels and with a dry or some- 
what fleshy, but never 
petal oid perianth ; and this 
passes over into very differ- 
ent forms by the suppres- 
sion of the floral-leaves, 
perianth and sporophylls 
(unisexual flowers are 
common), and by the close 
aggregatio7i of the flowers 
in the inflorescence. The 
flower is hypogynoui in 
every case. The inflores- 
cence is a spike which 
may be either single or 
branched, and has often a F ia .296.-Pias 8 ava(4ttaZ e a/um/ e m). 

thick and fleshy axis (a spadix). In Palms and Araceae it is en- 
veloped, at any rate prior to the opening of the flowers, by a very 
Targe floral-leaf, the spathe, which may be petaloid (Figs. 297, 301). 

The fruit is most frequently fleshy (berry, drupe) or a nut, never 
a capsule. The embryo is small, with large, fleshy endosperm 
(Fig. 299 C) ; very rarely the endosperm is wanting. 

The numerous plants 
belonging to this family 
are large, herbaceous or 
tree-like, and the leaves 
seldom have the usual 
Monocotyledonous form, 
i.e. linear Avith parallel 
venation, but most fre- 
quently have pinnate or 
palmate venation. 

Order 1. Palmae 
(Palms). The majority 
are trees with an un- 
branched, cylindrical stem. 

FIG. 296. A portion of the stem of Attale i 
having short internodes with persistent leaf-bases. 


and covered with leaf-scars or the bases of the leaf-stalks (Fig. 
296), and at the summit a rosette of large leaves closely packed 
together (Fig. 295). An. exception to this is found in Calamus 
(Cane, " Rotang "), whose thin, creeping or climbing stems have 
long internodes ; sparsely l branched is, e.g. the African Doum-palm 
(Hyphcene). Notwithstanding their often enormous stems the 
Palms have fibrous roots, like the bulbous Monocotyledons. The 
leaves are pinnate (Feather- palms, Fig. 298) or palmate (Fan- 
palms, Fig. 295) and often very large ; they have a well-developed 
petiole with an amplexicaul sheath, which is often more or 
less separated into a large number of fibres. In the bud the blade 
is entire but folded, as the leaf expands the lines of folding are 
torn, either those which are turned upwards (thus V V V V , e.g. 
Pritchardia, Livistona, Phoenix, Chamcarops) or those turned down- 
wards (thus A A A A, e.g. Cocos, Chamwdorea, Calamus). The 
inflorescence is usually lateral ; when, as in Sago-palm (J/t-- 
tro.rylon rumphii) or Talipot (Corypha umb mculifera) it is termi- 
nal, the plant is monocarpic, and dies after flowering; it is 
often a very large and branched spadix with numerous flowers 
either borne externally or embedded in it, and enclosed either in 
one woody, boat-shaped spathe (Fig. 297) or several, in 
the latter case one for each branch. The flowers are sessile or 
even embedded, regular, generally unisexual (monoecious or 
dioecious) with the usual diagram (Fig. 278) ; the perianth is in- 
conspicuous, green or yellow, persistent, and more or less leathery 
or fleshy. 6, rarely 3 or many stamens. The 3 carpels remain 
either distinct or form one, generally 3-locular, ovary. The style 
is short. There is one ovule in each carpel. Often during ripening 
2 carpels with their ovules are aborted. The fruit is a berry, 
drupe or nut, generally one-seeded, with a large horny or bony 
endosperm with hard thick- walled cells (e.g. Date-palm). In 
some (e.g. Cocoanut) it is thin-walled, soft, and oily; in several 
" ruminate." 

When //cnii i na tion commences iii the Cocoanut, Date, etc., the apex of the 
cotyledon remains in the seed and developesiuto a spongy mass to withdraw the 
eudosperm ; in the Cocoauut it attains a considerable size (Fig. 299 C) and 

1 [Although uubrauched stems are characteristic of the Palms, yet branched 
specimens are recorded from some eleven genera. The branches are developed 
from lateral buds, which in many instances only develope when the terminal 
bud bas been destroyed. A few Palms develope axillary branches at the base 
of the stem ; these form rhizomes, and give rise to clusters of aerial stems.] 


assumes the form of the fruit. The endosperm in the Cocoanut is hollow 
and the interior is filled with "milk." In the Date-palm and the Vegetable- 
ivory (Pliytelephas) the cell- walls of the hard endosperm serve as reserve 

I. PHCKNICEJ:. Phoenix (Date-palm) has pinnate leaves with 
channeled leaflets and dioecious flowers with 3 free carpels, of 

FIG. 297. Inflor- 
escence of a Palm 
with spathe. At the 
top c? -, at the base 
? -flowers. 

FlG. 2S. l.irittona austral if. 

which usually only one developes into a berry with membranous 
endocarp ; the large seed has a deep furrow on the inner side, and 
horny endosperm. 

2. SABALEJ:. These have fan-like leaves with channeled seg- 



ments ; flowers ^ or polygamous, rarely dioecious, with 3 separate 
or only slightly united carpels, all of which are sometimes 
developed into fruits (berry or drupe, with thin stone). 
Chamcej-ops, the Dwarf-palm. The pericarp is externally fleshy, 
internally more fibrous, and provided with a membranous inner 
layer. The endosperm is ruminate (that is, the testa is several 
times deeply folded into the endosperm). /S'aia/, Copernicia, 
Liristona (Fig. 298), Thriita.v, Corypha, Braliea, and others. 

FIG. 299. .-1 Longitudinal section of a Cocoanut (diminished), the inner layer only (the 
stone) not being divided .B End view of the stone, showing the sutures for the 3 carpels 
(a), and the 3 germ-pores ; the embryo emerges from the lowest one when germination 
begins. C Germinating; inside the stone is seen the hollow endosperm and the enlargir.^ 
cotj ledon. 

3. COCOLNEJS. With pinnate leaves. Monoecious inflorescence. 
The carpels are united into a 3-locular ovary. The fruit is most 
frequently 1-locular, only 1 of the loculi becoming developed, 
rarely 3-locular ; it is a drupe with a large, fibrous, external layer 
(mesocarp) and most frequently a very hard inner layer (endocarp, 
stone) which has 3 germ-pores, the 2 of these, however, which 
correspond to the suppressed loculi are closed; internal to the 
third lies the small embryo (Fig. 299). Endosperm containing 


abundance of oil. Cocos (the Cocoannt-palm), Attalea, Elms, 
Acrocomia, Bactris. 

4. LEPIDOCARYIX.E. The floral-leaves and flowers are borne in 
2 rows on the spadix. The carpels are united into one 3-locular 
ovary ; the fruit is coated by a layer of hard, shining, imbricate 
scales. The majority of the species are thorny, and climb by 
means of the thorny leaves. Some have fan-like (Mauritia), others 
pinnate leaves (RapMa, Calamus, Eugeissonia, Metro.rylon ; the 
stems of the latter die after the first flowering). 

5. BORASSIN/E. Large Fan-palms without thorns, with 3-locular ovary. 
Drupe with separate stones. Latania and Lodoicea have many stamens ; 
Hyplnfiie ; fiorassus (Palmyra-palm). 

6. ARECINE^E. The most numerous group. Feather-palms. Berry. Areca, 
Euterpe, Oreodoxa, Ceroxylon, Oiamtedorea, Geonoma, Caryota with bi-piuuate 

7. PHYTELEPHANTIN.E. Flowers with rudimentary perianth united in close 
capitula. Phytelephas (Vegetable-ivory). Nipa. 

DISTRIBUTION. About 1,100 species are known. In Europe only the Dwarf- 
palm (Chanuerops humilis) is wild (Western Mediterranean). The Date palm 
(Phoenix dactylifera) belongs to North Africa and West Asia. Other African 
genera are Hyphicne (Douna-palm) and Elms (E. guineensi*, Oil-palm). A 
large majority of the genera are found in South America and in the East 
Indies. The following are AMERICAN : Mauritia, Acrocomia, Bactris, Chaiiuc- 
dorea, Oreodo.ra, Euterpe, Attalea, etc. ASIATIC: Metroxylon, Calamus, Areca, 
JJorassH.s Lodoicea (" Double-cocoanut?," Seychelles) and others. The Cocoa- 
nut-palm has perhaps an American origin ; all the other species of the same 
genus being endemic in America ; it is the only Palm found on the coral islands 
of the Pacific Ocean, and is also the only one which is common to both hemi- 

USES. Palms belong to the most useful plants ; they contain no poison, and 
are of little medicinal interest, but are largely employed in the arts and manu- 
factures, the hard timber being adapted for many purposes on account of the 
hard tissue in which the vascular bundles are embedded. " Cane " is the 
stern of Calamus-species (from India). SAGO is obtained from the pith of 
Metroxylon rumphil (Sago-palm, Suuda-Is., Moluccas), Mauritia ftexuo*a, etc. 
Sugar-containing sap (" palm wine ") is obtained from the American Mauritia 
rinii'era and flexitosa, Borassus flaleUiformis (Asiatic Palmyra-palm), Arenga 
saccharifera, etc., by cutting off the young inflorescences, or by perforating the 
stem before the flowering (arrack is distilled from this). Vascular strands for 
the manufacture of mats and brushes, etc., are obtained from the outer cover- 
ing (mesocarp) of the Cocoanut, and from the detached leaf-sheaths of Attalea 
funifera (Brazil) (Fig. 296). WAX is yielded by the leaves of Coper nicia cerifera 
(caruaiiba-wax, Amazon region), and by the stem of Ceroxylon andicola 
(palm-wax, Andes) ; East Indian Dragon's blood is from the fruit of Calamus 
draco ; the young buds of ma-ny species, especially Euterpe, Cocos, Attalea, 
etc., are used as " cabbage." Palm-oil is obtained from the oily mesocarp 


of the plum-like fruits of Elceis yuineensis (W. Africa), and from the seeds, 
when it is largely used in the manufacture of soap. EDIBLE FRUITS from 
the Date-palm (Phcemx dactylifera, Arabia, Egypt, W. Africa), and the 
endosperm of the Cocoa-nut (Cocos nucifera}. The seeds and the unripe fruits 
of the Areca-palm (Areca cuteclni) are chewed with the leaves of the Betel- 
peper, principally in Asia. VEGETABLE IVORY from the hard endosperm of 
Pln/telephas mncrocarpa (S. America.) Many species are cultivated in the tropics 
as ornamental plants, hut in this country only Chamcerops htiniilis, Liciatouti 
australis and chinensis are generally grown. In addition to the few just 
mentioned, many others are of importance, but these are much the most 

Order 2. Cyclanthacese. This is a small order rein ted to the Palms (44 
species from Tropical America), with fan-like, folded leaves. The flowers are 
unisexual and arranged in whorls or close spirals on an uubranched spadix. 
Ovary unilocular, ovules numerous. To this belongs Carludouica palmata, 
whose leaves are us-cl for Panama hats. 

Order 3. Pandanaceae (Screw-pines) is another small order, forming a 
transition to the Aracpffi. The woody, (apparently) dichotomous stem is 
supported by a large number of aerial roots, which sometimes entirely support 
it when the lower portion of the stem has decayed. The leaves are closely 
crowded together, and arranged on the branches in three rows, which are often 
obliquely displaced, with the formation of three spiral lines ; they are, as in 
the Bromeliaceas, arnplexicaul, long, linear, the edge and lower midrib often pro- 
vided with thorus. The $ -flowers are borne in branched, the ^ in un- 
branched spadices or capitula, which resemble those of Sparganiuni, but have 
no floral-leaves. Perianth absent. The drupes or berries unite into multiple 
fruits. About 80 species in the islands of the Indian Ocean. Pundniiiia, 
Freycinetia. Fossils perhaps iu the chalk of the Harz. 

Order 4. Typhacese. The flowers are unisexual, monoecious, 
and borne on a cylindrical spike or globose capital urn ; 
inflorescences above, the ? below. The perianth consists of a 
definite number of scales (Sparganium), or in its place numerous 
irregularly-arranged hairs are found (Typha) ; in the (J -flower 
there are generally three stamens ; the gynceceum is formed of 1-2 
carpels with 1 prolonged style ; 1 pendulous ovule. The seeds are 
furnished with a seed-cover, which is cast off on germination. 
The few species (about 20) which belong to this order are marsh 
plants with cr/eeping rhizome (and hence grow in clusters) ; the 
leaves on the aerial shoots are borne in two rows, entire, very 
long and linear. 

Sparganium (Bur-reed). The flowers are borne in globose 
capitula ; the perianth distinct, generally consisting of 3 small 
scales ; pistil bi-carpellate. Drupe, dry and woody. The stalk of the 
lower $ capitula is sometimes unit- d with the main axis, and consequently the 
capitula are situated high above their subtending-leaf. 


TypJia (Bulrush, Reed-mace) has a long, cylindrical, brown 
spike, the lower portion bearing ? -flowers, and the upper <$ -flowers, 
which is divided into joints by alternate leaves. The $ -flowers 
have 1 carpel. The perianth is wanting, represented by a number 
of fine, irregularly-placed hairs; pistil unicarpellate. Fruit a nut. 

The two genera, according to some, are related to the 2nd order In both 
genera native species are found. The pollination is effected by the wind, and 
consequently the anthers project considerably, and the stigma is large and 
hairy. Typlia is protandrous, Sparganium protogynous. The small, fine 
hairs surrounding the nut of Typha. assist in its distribution by the wind. 
Fossil Typhas in the Tertiary. 

Order 5. Aracese (Arums). The flowers are small, and always 
borne without bracts or bracteoles on an unbranched, often very fleshy 
spike, which is enclosed by a spathe, often petaloid and coloured 
(Fig. 3 ',">!). The fruit is a Lerry. Outer integument of the seed 
fleshy. The leaves have generally sheath, stalk, and blade with 
distinctly reticulate venation ; they are chiefly cordate or sagittate 
(Fig 302), seldom long with parallel venation as in the other 
Monocotyledons (Acorus, Fig. 300). The Aracege are quite 
glabrous, generally perennial herbs with tubers or rhizomes. Many 
have latex. For the rest the structure of these plants varies ; for 
example, while some have a perianth, in others it is wanting ; in 
some the perianth-leaves are free, in others united ; some have 
hermaphrodite flowers, but the majority unisexual (monoecious) ; 
some have free, others united stamens ; the ovules are 
orthotropous, anatropous, or campylotropous, erect or pendulous ; 
the ovary is 1-many-locular ; some have seeds with endosperm, 
others without. In habit there are great differences. While some, e.g. Colo- 
casia (Fig. 302), have a thick, more or less upright stem, with leaf- scars, but not 
woody, others are climbers, epiphytic, and maintain themselves firmly by means 
of adventitious roots, on the stems and branches of trees, or even on steep rocks, 
e.g. Pliilodendron ; the cordate, peuninerved leaf is the most common (Fig. 
302), but various braucbed forms appear ; the pedate leaves of Helicophyllum, 
Draciincttlus, etc., are cymosely branched ; the leaves of filonstera delicioxd, 
perforated by tearing, should be noticed (the vascular bundles while ia the 
bud grow faster thun the tissue between them, causing the latter to be torn, 
and the leaf perforated). With regard to the anatomical structure, the 
presence or absence of latex, raphides, resin-passages, groups of mucilage- 
cells should be noted. Engler makes use of these anatomical peculiarities for 
a scientific arrangement of the order. 

A. OKONTIEJE, CALAMUS-GROUP. $ , hypogynous flowers of a 
completely formed monocotyledonous type (number in the whorls 
2, 3, or 4). Acorus (A. calamus, Sweet-flag) has a regular, 



3-merous, pentacyclic flower 
(Fig. 300 C, D). They are 
marsh-plants, with creeping 
rhizome, triangular stem, 
and long, sword-like leaves 
(Fig. 300 A) ; the inflor- 
escence is terminal, appa- 
rently lateral, being pushed 
to one side by the upright, 
sword-like spathe (Fig. 300 
r>).Anthnrium (Pr2 + 2, A2 + 2, 
G2); Pathos; Orontiiuii (unilocular 
ovary with one ovule), etc. 



FIG. 300. Acorus calamus: A habit (much 
reduced); B inflorescence; C a flower; D 
diagram ; E longitudinal section of an ovary j 
F an ovule. 

FIG. 301. 4nnu inoculation. The 
spathe (/i) in B is longitudinally 



B. CALLED. Flowers hypogynous, naked, ^ . - - Calla (G. 
palustris). All flowers in the spike are fertile, or the upper ones 
6-9 stamens ; ovary unilocular with many basal ovules. 


Marsh-plants with creeping rhizome and cordate leaves. Monstera, 
Rhaphidophora, etc. 

C. ARINE^. Flowers monoecious, naked, g -flowers on the upper, 

? on the lower part of the spadix. Arum (Fig. 301). The spadix 

terminates in a naked, club-like portion (A-) ; below this is a 

number of sessile bodies (rudimentary flowers), with broad bases 

FIG. 302. Colocasia Boryi. 

and prolonged, pointed tips (6); underneath these are the <$-. 
flowers (m),each consisting only of 3-4 short stamens, which eject 
vermiform pollen-masses through the terminal pores ; then follow, 
last of all, $ -flowers (/), each of which consists of one unilocular 
ovary, with several ovules. Perennial herbs, tuberous, with 

cordate leaves. Dractmculus ; Bwmm ; Arisarum ; Pinellin (Atherunis) ter- 
nata with leaves bearing 1-2 buds. Zantedeschin (ftliiapicn (Richanlia , Nile lily) ; 
$ , 2-3 stamens ; $ with 3 staminodes, 1-5-locular ovary (S. Africa.) In some 
genera sterile flowers are present between the $ and ? portions of the spadix 
(e.g. in Phllodendron) ; in Ambrosinia a lateral, wing-like broadening of the axis 
of the spadix divides the cavity of the spathe into two chambers, the anterior 
W. B. X 


containing one <j! , and the posterior 8-10 <? -flowers in two series ; in some the 
stamens in the single $ -flowers unite and form a columnar " synaudrium " 
(e.(j. in Die/enbacliia, Coloeasia, Alocasia, Caladium, Taccarum, Fyngonium). 
A remarkable spadix is found in Spatkicarpa; it is united for its entire length, 
on one side, with the spathe, and the flowers are arranged upon it in rows, the 
? to the outside, and the $ in the middle (Zostera has a similar one). Pista 
similarly deviates considerably, it is a floating water-plant, with hairy, round 
rosettes of leaves ; in it also the spathe and spalix are united ; at the base a 
$ -flower is borne, which consists of one unilocular ovary, and above several 
$ -flowers, each composed of two united stamens. 

BIOLOGY. The inflorescences are adapted for insect-pollination ; they are 
protogynous, since the viscous, almost sessile stigmas come to maturity and 
wither before the pollen, which is generally dehisced by apical pores, is shed ; 
some pollinate themselves freely by the pollen from the higher $ -flowers 
falling upon the $ -flowers below them, and in some it is conjectured that the 
pollination is effected by snails. The coloured spathe, and the naked end of 
the spadix (often coloured) of certain genera function as the coloured perianth 
in other orders ; during flowering a very powerful smell is often emitted. Arum 
maculatum is worthy of notice : small flies and midges creep down into the 
spathe, between the sterile flowers (Fig. 301 b), which are situated where the 
spathe is constricted, and pointing obliquely downwards prevent the escape 
of the insects ; in the meantime, the stigmas are in a condition to receive 
any pollen they may have brought with them ; after pollination the stigmas 
wither, and exude small drops of honey as a compensation to the flies for their 
imprisonment ; after this the anthers (HI) open and shed their pollen, the sterile 
flowers wither, and the insects are then able to escape, and enter and pollinate 
other inflorescences. In many, a rise of temperature and evolution of carbonic 
acid takes place during flowering ; a spadix may be raised as much as 30C. 
above the temperature of the surrounding air. Again, under certain conditions, 
many species absorb such large quantities of water by their roots that water is 
forced out in drops from the tip of the leaf ; this may often be observed in 

About 900 species in 100 genera. Home, the Tropics, especially S. America, 
India, and the Indian Islands, preferably in shady, damp forests growing 
as epiphytes upon trees, and on the banks of streams. Outside the Tropics 
few are found. Acorns calamus was introduced into Europe from Asia about 
300 years ago ; it, however, never sets any fruit, as the pollen is unfertile. In 
England Arum maciilatum is a very common plant; this and A. italicutu 
are the only native species. Coloeasia antiquorum comes from Polynesia 
and the Indian Islands, and also Alocasia macrorrhisa. Fossils in Cretaceous 
and Tertiary. 

USES. Many species have pungent, and even poisonous properties (e.g. 
Diefferibachia, Lagenandra, Aritm), which are easily removed by boiling or 
roasting; the rhizomes of many species of Caladium, Coloeasia (C. antiquorum, 
esculent a. etc.), are very rich in starch, and in the Tropics form an important 
source of food. An uncommon occurrence in the order is the highly aromatic 
rhizome of Acorus calamus; this contains calamus-oil and acoriu which are used 
in perfumery. Many are ornamental plants, e.g. Zantedeschia ccilriopica (South 


Africa), generally known as " Calla," and Momtera deliciosa many other 
species are grown in greenhouses. 

Order 6. Lemnaceae (Duck-weeds). These are tlie most 
reduced form of the Spadiciflorte. They are very small, free- 
swimming water-plants. The vegetative system resembles a small, 
leaf-like body (Fig. 303/-/), from which roots hang downwards; 
this branches by producing a new, similar leaf-like body, which 
springs from a pocket-like hollow (indicated by a dotted line in 
the figure) on each side of the older one, at its base (or only on 
one side). The branching is thus dichasial or helicoid (Fig. 303 A, where 
/i /')/">/"' indicate shoots of 1st, 2nd, 3rd, 4th generations respectively). The 
leaf-like bodies are, according to Hegehuaier, leaf-like stems, and thus Lemna 

FIG. 303. Lemna: A vegetative system ; ~B portion of a plant with flowers ; one stamen 
find tip of the carpel project ; the remaining portions being indicated by the dotted line. 

has no other leaves than the spathe and the sporophylls ; according to the in- 
vestigations of Engler they are sterns whose upper portion (above the " pocket ") 
is a leaf, which is not sharply separated from the underlying stem-portion. 
The inflorescence is a very much reduced Araceous-spadix, consisting in Lemna 
of 1 or 2 stamens of unequal length (1-stamened <J-tiowers), 1 unilocular 
carpel ( 9 -flower), and 1 thin spathe (B). [The same is found in Spirodela 
polyrrhiza, etc. , whose daughter-shoots begin in addition with 1 basal-leaf. 
Wolffia arrhiza, etc., have no roots, no spathe, and only I <$ -flower in the 
inflorescence (Engler).] On the germination of the seed a portion of the testa 
is thrown off as a lid, so that an exit is opened for the radicle. 19 species. In 
stagnant fresh water, both Temp, and Tropical. In Europe the species are 
Lemna minor, trisulca, gibba ; Spirodela polijrrlnza, and Wolffia arrhi~a, the 
smallest Flowering-plant. 


Family 4. Enantioblastse. 

The flowers in this family are hypngynous and have in part the 
general monocotyleclonous type with 5 trimerous whorls completely 
developed in a regular hermaphrodite flower, and in part the 
flowers so much reduced that the type is very difficult to trace. 
On the one hand the family is well developed and has capitate 
inflorescences (Eriocaulaceca) and on the other hand it is distinctly 
reduced (Centrolepidacece}. This family has taken its name from 
the fact that the ovule is not, as in the Liliiflorse and nearly all 
other Monocotyledons, anatropous, but ortliotropous, so that the 
embryo (/S/Vacmy) becomes placed at the end of the seed opposite 
(evavTibs) to the hilum. Large, mealy endosperm. The orders 
belonging to this family are by certain authors grouped with the 
Bromeliacece and Pontederiacece, etc., into one family, FARINOSE, 
so named on account of the mealy endosperm, the distinguishing 
character of the Liliiflorre then being that the endosperm is fleshy 
and horny. 

Order 1. Commelinaceae. The complete Liliaceous structure without great 
reductions in the number of whorls, but with generally few ovules iu each 
loculus of the ovary, is found in the Commeliiiaceje, an almost exclusively 
tropical order with about 317 species ; herbs, some of which are introduced into 
our gardens and greenhouses. The stems are nodose ; the leaves often clasp- 
ing ; the flowers are arranged in unipared scorpioid cymes, often so that they 
form a zig-zag series falling in the median line of the bracts, and after flowering 
they bend regularly to the right or left, outwards or inwards. They are more or 
less zygomorphic, particularly in the stamens, which in the same flower are of 
different forms or partially suppressed. The outer series of the perianth is 
sepaloid, the inner petaloid, generally violet or blue ; the filaments are some- 
times clothed with hairs formed of rows of bead-like cells (well known for 
showing protoplasmic movements). Fruit a trilocular ca2^siile with loculicidal 
dehiscence (generally few-seeded) ; in some a nut. The radicle is covered by 
an external, warty, projecting covering which is cast off on germination. 
The abundant raphides lie in elongated cells whose transverse walls they 
perforate. Commelina, Tradescantia, Tinnaiitia, Cyanotis, Dichorisandra. 

Order 2. Mayacaceae. This order is closely allied to the Coinrnelinaceae. 
7 species. American marsh- or water-plants. 

In many of the following orders of this family the flowers are united into 
compound inflorescences, with which is accompanied a reduction in the flower. 

Order 3. Xyridaceae (50 species). Marsh-plants with radical, often equit- 
ant leaves arranged in 2 rows, and short spikes on long (twisted) stalks. The 
flowers, as in the Comrnelinaceas, have sepals (which however are more chaffy) 
and petals, but the outer series of stamens is wanting. Capsule (generally 

Order 1. Rapateacese. Marsh-plants with radical leaves, usually in two 


rows, and several spikelets oil the summit of the main axis, clustered into a 
capitulum or unilateral spike. Each spikelet has numerous imbricate floral- 
leaves and one flower. 24 species. South America. 

Order 5. Eriocaulaceae. The " Composita among Monocotyledons," a 
tropical order. The flowers are borne in & capitulum surrounded by an involucre, 
very similar to that of the Composite. The flowers are very small, unisexual, 
$ and $ often mixed indiscriminately in the same capitulum ; they have the 
usual pentaeyclic structure ; the leaves of the inner perianth are often connate 
and more membranous than the outer ; in some the outer series of stamens 
are suppressed ; in each of the 3 loculi is one pendulous ovule. Capsule. The 
leaves are generally radical and grass-like. 335 species ; Eriocaiilun, Paepa- 
lanthus, etc., E. septangulare on the west coast of Scotland, and Ireland, and in 
North America. 

Order 6. Restiaceae. A small, especially S. African and S. Australian, 
xerophilous order (about 235 species), which is quite similar in habit to the 
Juucaceas and Cyperaceas. The leaves are often reduced to sheaths. The 
flowers are dioscious, the perianth as in JHIICHS, but the outer series of stamens 
suppressed. The ovary and fruit as in Eriocaulacese ; the ovary, however, may 
be unilocular, and the fruit a nut. lientio, etc. 

Order 7. Centrolepidaceae. These are the most reduced plants in the 
family ; small grass- or rush-like herbs. The flowers are very small, naked. 
Stamens 1-2, carpels l-oo. 32 species. Australia. Centrolepis (flowers 
generally with 1 stamen and 2-oo carpels). 

Family 5. Liliiflorae. 

The flower is constructed on the general monocotyledonous type^ 
with 5 alternating, 3-ruerous whorls (Fig. 278), but exceptions are 
found as in the Iridaceae (Fig. 279) by the suppression of the inner 
whorl of stamens ; in a few the position in relation to the bract 
differs from that represented in Fig. 278, and in some instead of the 
trimerous, di- or tetra-merous flowers are found (e.g. Majantliemum, 
Paris}. Flowers generally regular, hermaphrodite, with simple, 
petaloid, coloured perianth (except, for example, Bromeliacea?) ; 
ovary trilocular, generally with 2 ovules or 2 rows of ovules in the 
inner angle of each loculus (Fig. 304 C, D). Endosperm always 
present. A very natural family, of which some divisions in part 
overlap each other. The habit varies ; the leaves are haweverlong, 
entire, with parallel venation, except in Dioscoreaceee (Fig. 313). 

In the first orders of this family the flowers are hypogynous, and in the 
first of all the styles are free, and the capsule dehisces septicidally ; in the 
following the flowers are epigynous and in some reduced in number or uni- 
sexual ; capsule with loculicidal dehisceuce, or a berry. 

HYPOGYNOUS flowers : Colchicaceas, Liliacea?, Couvallariacese, Bromeliacea) (in 
part) . 



EPIGTNODS flowers: Amaryllidacese, Iridacese, Bromeliacere (in part), Dios- 

Order 1. Colchicaceae. The flower (Fig. 304.4) is $ , regular, 
hypogynous, trimerous in all five whorls (6 stamens) ; anthers 
usually extrorse. Gynceceum with 3 free styles (A, D) ; fruit a 
capsule ivith septicidal dehiscence (_Z) ; embryo very small (F). The 
underground stem is generally a corm or rhizome, seldom a bulb. 

A. VERATRE^E. Veratrum,; perennial herbs, stem tall with 
long interuodes and broad, folded leaves ; the flowers andro- 
monoecious, with free, widely opening perianth-leaves (Fig. 304 A}, 
and globular anthers; inflorescence a panicle. Zygadenus, 
Melanthium, Schcenocaulon, Uvularia, Tri'cyrtis. 

B. TOFIELDIEJ;. Nartheciiim and Tojieldia have leaves alter- 
nate (arranged in two rows), sword-like and borne in rosettes ; 
racemes or spikes. Narthecium forms an exception to the order by 
having a simple style and fruit with loculicidal dehiscence ; 

FIG. 304. Veratrum : A flower; B stamen ; C transverse section of ovary ; 7) gynoeceurn, 
with one carpel bisected longitudinally, and the third removed ; E fruit after dehiscence ; 
f longitudinal section of a seed. 

Tofieldia by the introrse anthers. In this they are related to the 
Liliaceaa. Narthecium has poisonous properties, like many other 

C. COLCHICE.E. Colchicum (Autumn Crocus) ; perennial herbs, 
with a long, funnel-shaped, gamophyllous perianth, and introrse 
anthers. The flowers of C. autumnale spring up immediately 
from the underground stem, which is in reality a corm formed of 
one internode. Colchicum autumnale flowers iri autumn without leaves ; in 
spring the radical foliage-leaves appear simultaneously with the fruit. The 
flower is protogynous, and is pollinated hy insects (humble-bees, etc.) which 
seek the honey secreted by the free part of the stamen a little way down the 
tube. The length of the tube protects the fruit, and not, as in other cases, the 
nectary. Bulbocodinm and Merendera have unguiculate perianth-leaves, free, 
but closing together like a tube. 

175 species ; chiefly in North America and South Africa. Tofieldia is an 


Arctic plant. The order is rich in pungent, poisonous alkaloids (veratrin, 
colchicin, etc.). OFFICINAL; the seeds of Colchicum autumnale (Europe) and 
Sclifenocaulon officinale (Mexico), and the rhizome of Veratrum album (mountains 
of Central Europe). 

Order 2. Liliacese (Lilies). Flowers as in the Colchicacete 
but with introrse anthers ; ovary free, 3-locular, with single style ; 
capsule 3-locular with loculicidal dehiscence. The majority are 


FJG. 305. Colchicum autumnale. A Conn seen from the front : fe corm ; s's" scale-leaves 
embracing the tiower-stalk ; tr/i base of flower-stalk with roots (IP). B Longitudinal section 
of corm and flower-stalk : hh brown membrane surrounding the underground portion of 
the plant ; st flower- and leaf-stalk of previous year, the swollen basal portion forming the 
reservoir of reserve material. The new plant is a lateral shoot from the base of the corm 
(I;) and has the following parts : the base bearing the roots (to), the central part (fc') which 
becomes the corm in the next year, the axis bearing the scale-leaves (s', s"), the foliage- 
leaves (I, I"'), and the flowers (b, V) which are borne iu the axils of the uppermost foliage- 


herbs with bnlbs ; the inflorescence is terminal. In many species repro- 
duction takes place by means of bulbils (small bulbs) formed in the axils of 
the foliage-leaves (e.g. Lilittm bulbiferiim, htucifolium, etc., Gagea lancifolia, 
etc.), or in the bracts of the inflorescence (many species of AUiitm) ; in many 
species several buds are developed as bulbs in the axils of the bulb-scales them- 
selves (accessory buds arising close together), and in some the formation of 
buds is common on the leaves. 

A. TULIPEJ:, TULIP GROUP. Bulbs. The aerial, elongated stem 
bears the foliage-leaves. Flowers few but generally large, with 
free perianth-leaves. Tulipa ; style absent, no honey ; flowers 
generally solitary, erect. Fritillaria perianth campanulate with 
a round or oblong nectary at the base of each perianth-leaf. 
Lilium; perianth widely open, generally turned back with a 
covered nectary-groove in the centre of each segment. Anthers 
versatile. -Lloydia ; Erythroiiium. 

B. HVACIXTHEJE, HYACINTH GKOUP. Bulbs. Leaves radical ; aerial 
stem leafless with raceme or spike. In some the perianth-segments 
are free, in others united. Honey is produced often in glands or 
in the septa of the ovary (septal glands). Ornithogalum has a 
leafy stem ; Scilla ; Eucomis has a tuft of floral-leaves above the 
raceme; Agrapliis; Hyacinthus ; Puschkinia ; CMonodoxa; Muscari ; 
Velfheimia ; Urginea. 

C. ALLIED, ONION GROUP. Generally bulbs. Leaves radical. 
Stem leafless with a compound umbellate or capitate inflorescence 
of unipared helicoid cymes, which before flowering are sur- 
rounded by two broad involucral leaves. Allium. Filaments often 
petaloid and bidentate ; iu many species bulbils are found in the inflorescence. 
Some species have flat leaves: A. satirum, Garlic; A. pornun, Leek; A. 
umiuum; others have round, hollosv leaves: A. cepa, Onion; A. fistulas um, 
Winter Onion; A. ascalonicitm, Eschalot; A. scJuenoprasunt, Chive. Gagea ; 
honey is secreted at the base of the perianth, no special nectary; 
inflorescence few-flowered Agapanthus ; Tnteleia. 

D. ANTHEBICEJE. Rhizome; raceme; the leaves not fleshy and thick. 
Anthericnm ; Aaphodelns ; Bulbine ; ChloropJnjtioii ; Jioicica has an almost leaf- 
less stem with curved, climbing branches. 

E. ALOINE.E, ALOES. Stem generally aerial and tree like, bearing on its sum- 
mit thick, fleshy leaves, often with a thorny edge (Fig. 306). Eacerne branched 
or unbranched. Aloe ; Gasteria ; Yucca (has secondary thickening, p. 274). 

F. HEHEROCALLIDE.E. Phorwium, (Pli. tenax. New Zealand Flax) ; Fttnckia 
(Hos(a); Hemcrocallis. 

At this point the following are best placed: Apliyllanthi-s (A. monspeliensia) ; 
Xanthorrluea (Black-boy); Xerotes ; Lomandm ; Kingia ; the very membranous, 
dry perianth of the last resembles that of the Juncacere, and also there are only 
1-few ovules in the loculi. 


POLLINATION by insects. Honey in some is produced on the perianth (see 
Tulipea?), in others by glands on the carpels (in the septa and parietal placenta?, 
septal glands): Hyacinthus, Allium, Anthcricum, Asphodelus, Yucca, Fitnckia, 
Hcincrucallis, etc. Some ^Z^am-species ate protandrous. Fritillaria is visited 
by bees, Lilium martagon by moths, L. bulb if e rum by butterflies, Pliormiuin 
(New Zealand) by honey-birds. 

FIG. 306. Aloe. 

About 1,580 species ; rare in cold climates ; their home is in sunny plains 
with firm, hard soil, and warm or mild climate, particularly in the Old World 
(S. Africa; As. Steppes; Mediterranean) ; at the commencement of spring the 
flowers appear in great profusion, and after the course of a few weeks disappear ; 
during the hot season their life lies dormant in the bulb, hidden underground. 



The woody species are tropical. The majority of the introduced Liliaceae 
( fr'ritillaria imperialis, Crown-imperial; Lilium caiididitin; Tulipa gesnenana; 
Hyacinth ; Afrwcart-species ; Sc ilia-species ; Ornithogalum nutans; Hemerocallis 
jnlra andflava; Aspltodelus liitens aud albus) come from the Mediterranean and 
W.Asia; friinckia, from China and Japan; several Lilies from Japan and the 
Himalayas ; Agapanthus from the Cape ; Allium saticitm is a native of the 
Kerghis Steppes ; A. cepa from Persia (?) ; A. ascaloiiicum is not known 
wild (according to others a native of Asia Minor), perhaps a form of A. cepa ; 
A. ichosnoprasum from the N. temp, region. 

Many bulbs have pungent properties ; many Onions are used as culinary plants 
The bast fibres of Phurmium tenax (New Zealand Flax) are used technically. 
Dyes are obtained from the Aloe ; gum for varnish from the stem of Xanthorrhnea 
h-istiln and australe. OFFICINAL; "Aloes," the dried sap of S. African species 
of Aloe. (A. Africana, A.ferox, etc.) ; the bulb known as "Squills" from Urginea 
(Scilla) niaiitima (Mediterranean). 

Order 3. Convallariaceae. This order differs from the 
Liliaceae in having the fruit a berry (Fig. 308) and in never being 
bullions ; the seeds are less numerous. 


307) and normal foliage-leaves. Polygonatum : rhizome creeping ; 

aerial shoot leafy, bearing the 
flowers in racemes in the axils of 
the foliage-leaves ; perianth tubu- 
lar. P. multiftorum (Solomon's 
seal), P. officinale, etc. Majanthe- 
mum : flower 2-merous ; perianth 
almost polyphyllous, spreading. 
Smilacina. Streptopus (S. amplexi- 
fulius ; the flowers or inflorescence 
unite with the entire succeeding 
internode). Convallaria (1 species 
C. majalis, Lily of the valley) ; flowers in terminal racemes ; 2 basal 
foliage-leaves ; perianth globose, bell-shaped. lleineckea earned 
(Japan, China) in gardens. Paris (P. quadrifolia, Herb- Paris) ; 
flowers solitary, terminal, 4-merous, polyphyllous ; styles 4, free 
(approaching the Colchicacea? ; it is also poisonous) ; a whorl of 
4 (-more) 3-nerved, reticulate leaves on each shoot. Ornamental 
plants: species of Trillium, Aspidistra elatior (Japan). 

B. ASPAKAGEJ;, ASPARAGUS GKODP. Scale-like leaves and green 
assimilating branches. Asparagus : horizontal rhizome. The aerial 
shoots are very richly branched ; the numerous needle-like bodies 
upon the plant are leafless shoots, which are crowded together in 
double scorpioid cymes in the axils of the scale-leaves ; the two first 

FIG. 307. Rhizome of P.<Z!/goi!iiti<iii 
multijlorum : a bud ; b shoot ; c d scars 
left by shoots of previous years. 


lateral axes, placed outside to the left and right, generally bear 

FIG. 308. Smilax pseudosuphilitica : A shoot of male plant ; C <J -flower ; D berry, almost 
ripe; E the same in longitudinal section. B Smilax syphilitica : portion of branch with base 
of leaf and tendrils. 


flowers. Polygamous. Puscus (Butcher's broorn) is a S. European shrub 
with leaf-like, ovoid or elliptical shoots (phylloclades) which are borne in the 
axils of scale-like leaves, and bear flowers on the central line. Dioecious. 
Stamens 3, united, anthers extrorse. Semele androgyna bears its flowers on 
the edge of the flat shoot. 

C. SMILACEJ;. Smilax (Sarsaparilla) (Fig. 308) ; climbing 
shrubs with the leaf-sheath produced into tendrils. The leaves 
have 3-5 strong nerves proceeding from the base, and are reticulate. 
Orthotropous or semi-anatropous ovules. Dioecious (Fig. 308 C,E). 

D. DRAC^NE^E. Fruit in some a berry, in others a capsule. The stem of 
DRACAENA, when old, has the appearance of being dichotoruously branched ; it 
has the power of increase in thickness, and may become enormously thick. The 
Dragon-tree of Teueriffe, measured by Humboldt, attained a circumference of 
14 in. and a height of 22 m. ; the leaves are large, linear or linear-lanceolate. 
Cordylitie (East Asia), various species in gardens and greenhouses (Yucca is 
closely allied). Astclia. 

POLLINATION. Pans quadrifolia and Concallaria majalis have no honey, 
and are chiefly visited by pollen-collecting bees (in the absence of insect visits 
pelf-pollination takes place) ; Polyyonatuin mult'itiorum has honey secreted by 
septal glands and protected by the base of the tubular perianth ; it is pollinated 
by humble-bees, etc. Asparagus vfftcinalis has small, polygamous, greenish, 
honey-bearing flowers ; the $ -flower is almost twice as large as the 9 ; both 
have rudiments of the opposite sex. 

About 555 species ; especially from N. America, Europe, and Central Asia. 

OFFICIAL : " Dragons' -blood," a red resinous juice from the stem of 
Dracaena and the roots of some Central American species of Smilax. The 
tuberous stems of the Eastern Asiatic Smilax glubra are officinal. The flowers 
of Coitrallaria majalis have been lately used as a substitute for Digitalis. 
Pungent, poisonous properties are possessed by Paris. None of the species are 
used as food, except the young annual shoots of A^ai'ayua officinalis, a shore- 
plant which is used as a vegetable. 

Order 4. Pontederiaceae. Flowers generally zygomorphic, hypogynous, $, 
with handsome, white or violet, petaloid perianth which forms a tube at its base. 
The stamens are inserted at different heights in the perianth-tube, and are re- 
duced to three (in Heteraiitheia seldom to one). In some the ovary is trilocular 
with GO ovules (Eichliornia), in others reduced to one loculus with one ovule 
(Pontederia). Fruit a capsule or nut. Embryo as long as the abundant, mealy 
endosperm. Tropical water-plants (22 species) with peculiar sympodial branch- 
ing, nearly the same as in Zo-itera. Spikes without floral-leaves. Many inter- 
cellular spaces in the stem and leaf. In greenhouses: Eichliornia a z urea, E. 
i-rassipex (both from tropical and sub tropical S. America) ; the latter has 
swollen petioles which serve as floats and enable it to float freely on the water, 
sending down its roots into the mud. Hete'rantheia reniformix, H. zosterifoiia, 
Pontederia cordata. 

Order 5. Amaryllidacese (Narcissi). The flower is epi- 
gynous, otherwise exactly the same as in the Liliacea? (6 stamens). 



The majority, like these, are also perennial herbs with bulbs and 
scapes. The fruit and the other characters as in the Liliacese. 
The external appearance is, however-, very different. 

A. AMARYLLE.E have bulbs and the leaves generally arranged 
in two rows ; the flowers are borne singly or in umbel-like in- 
florescences on lateral scapes, while the main axis of the bulb is 
unlimited. Beneath the inflorescence is an involucre (Fig. 309). 
Galanthus, Snowdrop, has a polyphyllous perianth without coi'ona; 
the three inner perianth-leaves are emarginate and shorter than 

FIG. 309. Pancratium canbceum. 

the outer ; the anthers dehisce apically. Leucnjnm differs in haviug 
the perianth-leaves equal in length. Amaryllis has a funnel-shaped perianth, 
entirely or nearly polyphyllous, but somewhat zygomorphic. Criniim ; 
H(cmantlms ; Clivia. Narcissus has a tubular corona, a ligular struc- 
ture arising from the perianth-tube exterior to the outer stamens. 
In Pancratium (Fig. 309) the corona is united with the filaments which appear 
to spring from its edge. Eticharis amazonica. 

B. HyroxiDEJE. The leaves, which are grass-like, dry, folded, and in some 
hairy, spring from a rhizome, generally with a divergence of J /s. Flowers small, 
perianth polyphyllous, persistent, on which account perhaps the Hypoxideae may 


be considered as the least altered type. The chief characteristic is that the 
embryo is separated from the hiluru. Hypoxis ; Curcnligo (C. recurrata, a 
favourite ornamental plant; S.E. Asia). 

C. ALSTKOSMEKIE^E. (Alsti'uemeria, Bmnarea) ; stems long, leafy, often climb- 

D. VELLOSIEJE (Vellosia, Barlaccnia) stem woody, usually rlichotomously 
branched, with terminal, single flowers ; it bears numerous aerial roots which 
pierce the leaves and surround the stem. Stamens often (by splitting) 6-18. 
High table-lands of S. America and S. Africa. 

E. AGAVES. Very similar to the Bromeliacese both in their distribution 
(nearly all American) and in external appearance. They appear as gigantic bul- 
bous plants with perennial, aerial, generally short stem, and perennial, large, 
lanceolate or linear, stiff, thick, and often thorny leaves, which form a large 
rosette; after the course of several (8-20) years the terminal inflorescence is 
developed, which is 10-12 m. high, paniculate, and freely branched. Before 
the inflorescence expands, a large quantity of sugar-containing sap is collected 
from A. americaiia by removing the terminal bud; this on distillation yields 
" pulque," the national drink of Mexico. After flowering the entire shoot dies, 
but the subterranean lateral shoots survive and reproduce the plant. A nave 
aniericana, etc. ; Fourcroya ; Polianthcs titberosa (Tuberose ; Central America). 

DISTRIBUTION. The 650 species are chiefly natives of S. Africa and S. 
America. Clivia, Hminaiitlms, Amaryllis are from the Cape; Narcissus from S. 
Europe, whence many species have been introduced ; Galantlms and Leucojiiin 
are especially from S. and Central Europe, and from the Caucasus. 

USES, few, except as ornamental plants : Galanthus nivalis ; Leucojuin : Nar- 
cissus pseudonarcitsus, N. poeticits, N. jonquillti, N. tazetta, etc. ; Amaryllis, 
Altra'ineria, Enchant, Criuuni, Ta/lota, etc. The \ ascular bundles of the various 
species of Agave (Agave rigida, var. sisalana, sisal hemp,) are used for cordage, etc. 

Order 6. Bromeliaceae. The flowers are hypogynous, 
epigynous or semi-epigynons ; the perianth is divided into calyx 
and corolla; stamens 6. The fruit is a capsule or berry with 
many seeds. Endosperm mealy, embryo small, at the edge of the 
endosperm, but not enclosed by it. 

Perennial herbs with a very characteristic appearance (Fig. 310) ; 
the stem is most often short, thick, and crowned by a rosette of 
many leaves, which are long, often very narrow, leathery, stiff, 
and with a spiny edge ; they are usually channeled, completely 
closing round each other, with their edges forming a tightly closed 
hollow, in which generally water is collected (this among other 
things insulates the inflorescence and thus prevents the access of 
creeping insects, such as ants). The presence of numerous stellate, 
water-containing hairs often gives the leaves a grey appearance, 
and the layers of cells beneath the upper epidermis of the 
lamina form an " aqueous tissue," which serves as a protection 
against the rays of the sun and regulates the evapoi'ation. The 



FIG. 310. Aechinta miniata. 

stomata are often situated in 
farrows on the under-side of 
the leaf, and hence cause a 
striped appearance. They are 
all American (525 species), es- 
pecially from S. America, where 
they live partly as epiphytes on 
trees, partly in the clefts of rocks, 
often on the steepest slopes, to 
which they firmly attach them- 
selves by aerial roots; some are 
terrestrial. The stem is seldom 
tree-like or many metres in 
height (Puya, in Chili ; Hechtia, 
in Mexico). The inflorescence is 
a terminal spike, raceme, or 
panicle, often with large and 
brightly-coloured floral-leaves. 
The flowers are without scent. 

FIG. 311. Multiple-fruit of Ananassa sativa* 


The seeds, in the species whose fruit is a capsule, are often pro- 
vided with wings (hairs, expansions, etc). Ananassa satiua, Pine- 
apple (W. Indies, Central America) is cultivated for the sake of 
its juicy, aromatic fruits, which coalesce with their fleshy bracts 
and form a large spike-like fruit-cluster (multiple-fruits, 1 Fig 311) 
bearing on its apex a leafy shoot, which may be used as a cutting. 
Seeds very rarely developed. Tillandsia (T. usneoides is a fila- 
mentous, richly branched, rootless epiphyte hanging in masses 
from trees; Trop. Am.), Aechmea, Sillbergia, Pitcairnia, etc. 

USES. The leaves of the Pine-apple, in its native country, are used for the 
manufacture of cloth. 

Order 7. Haemodoraceae. 120 species ; in all parts of the world except 
Europe ; perennial, often tornentose and resembling the Bromeliacese, Iridaceas 
and Amaryllidaceffi. Hcemodoriini (Australia). To this order belong Ophio- 
pogon, Peliosanthes, Sanscviera, and others. 

Order 8. The Iridacese have epigynous, hermaphrodite flowers 
with petaloid perianth as in the Amaryllidacea?, but the interior ichorl 
of stamens is entirely suppressed, and the 3 developed outer stamens 
have extrorse anthers (Fig. 279) ; there is 1 style with 3 large, gene- 
ral \yinore or less leaf -I ike branches bearing the stigmas. Ovary and 
capsule as in the Amaryllidaceaa and Liliacete. Perennial herbs ; 
bnlbs are rarely found, but horizontal rhizomes, corms, etc., take 
their place. The leaves are (except Crocus} as in the Jn'.s, tn-<>- 
rotved, equitant and sword-like. Flowers or inflorescences terminal. 

The Iris (Flag) has a horizontal rhizome. The flowers are borne 
in the leaf-axils in fan-like inflorescences (rhipidium). The 
branches of the style are large and petalaid ; on their under surface 
may be seen a small projecting shelf (Fig. 312 a) having on its 
upper surface the stigmatic hairs. Beneath the branches of the 
style are 3 well protected stamens, and immediately outside these 
the external perianth-leaves. The honey is secreted in the perianth-tube, 
and the insects, endeavouring to obtain it through the narrow passages at the 
base of the stamens, settle upon the outer perianth-leaves, which are bent back- 
wards and often very hairy along their central line. The insects then rub 
their backs on the anthers just above them, beneath the branches of the style ; 
they readily deposit the pollen on the stigma of another flower as they enter it, 
but cannot do so in withdrawing, since the stigma is pushed back, and self- 
fertilisati'in is thus avoided. The stylar branches lie close to the outer 
perianth-leaves, which are just beneath them, or separated by a distance of only 
6-10 mm. ; the first form of flower is adapted for Rlnnqia ro*truta, the 
latter for bees. Crocus has vertical, tuberous, underground stems 

1 The aggregation of the fruits of several distinct flowers into one mass. 



surrounded'by the leaf-sheaths (corms), and terminal flowers; the 
linear leaves are not equitant, but have two longitudinal furrows 
on the under side. The perianth is gamophyllous and funnel- 
shaped. The stylar branches (stigmas) are fleshy, rolled together 
in the shape of a horn, and split along the edge. Gladiolus has 
corms like the Crocus ; spikes with slightly zygomorphic, almost 
bilabiate flowers, most frequently turning to one side. Position 
of the leaves as in the Ir\s.Dii>hnrhi>n<i has 2 fertile and 1 barren 
stamen ; Hennodactylns has a unilocular ovary with 3 parietal placentae. 
Cypella and Tigridia have bulbs. 


FIG. 312. Iris pseudacorus. One external and two internal perianth-leaves, and one 
of the stylar-branches have been removed, y The outer, i the inner perianth-leaves ; 
g stylar-branch ; a stigma; s anther. The ovary is seen in longitudinal section. 

770 species ; chiefly in the countries round the Mediterranean, and in Africa, 
especially the Cape ("ladiolus, Ferraria, Moraa, GaJaxia, Sparaxis, Antholt/za, 
Tritonia, I.ria, etc.), Australia and Tropical America (Sixyrinchinm, Tirjridia, 
Cipiiru, Cijpella, etc). A great number are ornamental plants : the cultivated 
Crocus-species are from the South of Europe and Asia ; Gladiolus cominu-iis 
from S. Europe ; the other species principally from S. Africa. The native 
species of Iris are 7. pseudacorus (yellow) and I. feetidissima. 

OFFICINAL : the stigmas of Crocus aativus (Oriental, cultivated in France, 
Spain, Italy, and Austria), used as a colouring matter, saffron ; the rhizomes 
of the S. European Irisjiorentina, pallida, and germanica (" Orris-root"). 
W. B. v 



Order 9. Dioscoreaceae. Perennial herbs with fleshy, often 
very large tuberous rhizomes (or roots) ; twining stems ; leaves 
stalked, often arrow- or heart-shaped, lobed, palminerved and finely 
reticulate as in the Dicotyledons (Fig. 313). The flower is 
diclinous (most frequently dioecious), regular, epigynous, small, and 







;: s 

<B a) 

03 -. 
*0 'S 

te ui 

.. m 




of a r/reenish colour, but otherwise typical (Pr3 + 3, and A3 + 3, 
or CI3) ; in most instances 2 ovules are placed one above the 
other in each loculus. The inflorescence is a spike or raceme, some- 
times richly branched and paniculate. The order approaches most 
nearly to the Amaryllidacese. 


Tamus (Bryony) has a berry, Dioscorea (Yam) a thin-walled, 
3-edged or 3- winged capsule (Fig. 313). Both have subterranean 
or aerial tubers ; the Yam very often also developes tubers in the 
axils of the foliage-leaves ; tuberous roots are said to occur in 
D. batatas. The tubers of many species of Yams (D. batatas from 
China and Japan, D. alata, South Sea Islands and India, D. 
bulbifera) are a very important source of food in the Tropics, 
especially the first-named. Testudinaria ; Rajania. The tuberous 
stein of Tamus communis and Testudinaria elephantipes, and some species of 
Dioscorea is formed from one single internode (epicotyl), and the aerial shoots 
are developed from adventitious buds ; in T. elephantipes the stem is aerial, and 
covered with thick scales of cork, regularly arranged, and separated by grooves. 
Tropical order (167 species) ; 2 species (Tamns communis and Bordered 
pyrenaica) in Europe. 

Family 6. Scitamineae. 

The flowers belong to the ordinary monocotyledonous type. 
They are hermaphrodite, epigynous, and have either a petaloid 
perianth, or calyx and corolla ; they are, however, zygomorphic 
or unsymmetrical, and of the stamens most frequently only one 
is completely developed, the others being generally represented by 
petaloid staminodes. The ovary has 3 loculi, more rarely it is 
unilocular with the suppression of 2 loculi. Endosperm is 
absent (except Zingiberacece) ; but, on the other hand, there is a 
large perispenn. To this family belong large, glabrous, especially 
perennial herbs with rhizomes ; leaves large, distinctly divided 
into sheath, stalk, and blade, the latter being moi-e or less 
elliptical or lanceolate, entire, with pinnate venation, and always 
with a very well-pronounced midrib, gradually tapering towards 
the apex, and giving off numerous branches, which run outwards, 
towards the margin, at a lai'ger or smaller angle ; these lateral 
veins are closely packed, and parallel, but with only weak, con- 
necting branches between them ; the leaves, therefore, are easily 
torn pinnately (Figs. 314, 317). The leaf-sheaths close tightly 
round each other and form a false stem. 

This very natural family comprises orders closely connected 
with each other, but is not itself nearly allied to any other family. 
First in the series stands : 

Order 1. Musacese. The petaloid perianth is strongly 
zygomorphic, the anterior leaf being very large (a kind of 
" labellum "), the posterior one small ; only the posterior stamen 
is wanting, or is rudimentary, the other five are developed, and 



have quad ril ocular anthers ; ovary, 3-locular. Seed with straight 
embryo in mealy perisperm. 

The best-known genus is Musa, the Banana (Fig. 314). From 
the short rhizome arise enormously large, spirally-placed leaves, 
whose sheaths envelope one another, and form an apparently aerial 

FIG. 3H. Two ifMsa-species. 

stem, several metres in height. The inflorescence is a terminal 
spike with floral-leaves placed spirally, and sometimes magnifi- 
cently coloured ; in the axils of each of these several flowers are 
situated in two transverse rows (accessory buds) ; the lowest 
flowers in the inflorescence are ? , the central ones , the upper 



ones <$ , so that fruits are only found in the lower region of the 
inflorescence, the remaining portion persisting as a naked axis 
after the floral-leaves and flowers have fallen off ; the inflorescence 
terminates in an ovoid bud formed by the flowers which have not 
yet opened (Fig. 314, the left-hand figure). The perianth-leaves 
are united (except the posterior one). The fruit (known as 
a " Banana ") is a berry, having the form of a smooth, short, three- 
cornered Cucumber (as much as 30 cm. in length) ; inside the 
tough skin is found a farinaceous, aromatic pulp. No seed is 
developed in the cultivated species. Several MuM<-species are culti- 
vated in the Tropics for the sake of the fruit (M. -paranisiaca, AJ. 
sajnentttm) ; for the fibrovascular bundles, HI. text-ills (Manilla Hemp). Their 
home is, no doubt, the Tropics of the Old World ; they were introduced into 
America before the arrival of Europeans. Musa ensete has dry, leathery fruits; 
an ornamental plant. 

In 3!nsa the barren, posterior stamen belongs to the inner whorl ; and also in 
Rtrelitzia and Ravenala ; the latter may have all 6 stamens developed. In 
Heliconia, on the contrary, it belongs to the outer whorl; in Heliconia the 
perianth-leaves are differently arranged, and there is only one ovule in each 
loculus. The three latter genera have dry fruits and leaves arranged in two 
rows. In the "Travellers' Palm" (Ravenala madagascariensis) the foliage- 
leaves form an enormous fan. Tropical ; about 50 species. 

The order may be divided as follows: 1. Museas : Musa, 
Ravenala, Strelitzia in the Old World. 2. Heliconiae : Heliconia in 
the New World. 

Order 2. Zingiberaceae. Perianth most frequently divided 
into calyx and corolla. Calyx 
gamosepalous. Only 1 fertile 
stamen (the posterior, Fig. 315, 
belonging to the inner whorl) 
with quadrilocular anther, 
which encloses the style in a 
furrow ; the 2 stamens in the 
outer whorl are staminodes, 
the median one (the anterior) 
is wanting. The 2 lateral 
staminodes of the inner whorl 
form the " labellum " (Fig. 
315 lab), which usually is the 
largest segment of the flower, 
and is often bilobed. Ovules 
many. The fruit in some is 
a leathery, 3-valved capsule, 

FIG. 315. Diagram of a Zingiberaceous 
flower (Kiimpferia ovalifolia) : b bract; v brae- 
teole ; k calyx ; p 1 , p 2 , p 3 the petals ; ssf, 
lateral staminodes ("-wings''); lab labellum 
(formed of two staminodes); st the fertile 
stamen; * position of suppressed stamen. 
The ovary is in the centre of the diagram. 



with loculicidal dehiscence ; iu others it is more or less berry-like 
and indehiscent, or irregularly dehiscent. Straight embryo. The 
aerial stem is seldom developed to any extent, and the inflor- 
escences, which are (compound) spikes or racemes, often with 
coloured floral-leaves, spring in some (e.g. Zingiber nfficinale) 
directly from the rhizome. The leaves are arranged in two rows. 

-The ovary in a few instances (Globba and others) is unilocular, 

with 3 parietal placentae. 

They are perennial herbs with fleshy and tuberous rhizome*, which are used 
as condiments and in medicine on account of their pungent and aromatic 
properties and also for starch, dyes, etc. OFFICINAL : rhizomes of Zitigiber 
officinale (Ginger, unknown wild, but cultivated generally in the Tropics), of 
Curcuma longa (Turmeric, a dye, E. India) and C. zedoaria, of C. angustifolia 
and others (as E. India Arrowroot), of Alpinia officinarum, China (galangal). 
"Preserved Ginger" from Alpinia galanga. Similar aromatic materials (vola- 
tile oils) are present also, for example, in the fruits ; Cardamom fruits and seeds 
(from Elettaria cardamomum, China, seldom from E. major). 

315 species ; Tropics, preponderating in the Eastern Hemisphere, India, and 
especially S. Asia, whence all the aromatic species originate ; they are now 
commonly cultivated in the Tropics. Some are ornamental plants in green- 
houses, e.g. Hedychium, Costtis, etc. Globba (with axillary buds in the in- 
florescence, as in Ficarin), Renealmia, Kdmpferia. 

Order 3. Cannaceae. 

FIG. 316. Flower of Canna : / 
ovary ; pa calyx ; pi corolla ; I la- 
bellum ; st stamens ; an anther ; 
tj stigma ; a and p stamin odes. 

American herbs without aromatic pro- 
perties. Flowers asymmetric (Fig. 
316). Calyx polysepalous. The sta- 
mens are petaloid (Fig. 316 st) and 
barren with the exception of one (the 
posterior), which bears on one of its 
edges a bilocular anther ; another, 
which is especially large and coloured, 
is termed the labellum. The style is 
compressed and leaf-like, with a 
small stigma at the apex. Ovules 
numerous in the 3 loculi. The cap- 
sule is furnished with warts or soft 
prickles. Embryo straight. 

Canna (30 species ; Trop. Am.). 
The inflorescence is a terminal spike 
with 2-flowered unipared scorpioid 
cymes in the axils of the floral-leaves. 
Ornamental plants : Canna indica, etc. 

The diagram of the andrcecium of 


the Cannaceee and Marantaceas may be represented in the follow- 
ing manner (calyx, corolla and gynceceum being omitted) : 


w w 


w lab 





wi c 


ic The lateral staminodes, " wings ; " st fertile stamen ; * the suppressed stamen ; lab 
labsllutn ; c hood ; u'i inner- wing. 

The labellum of the Cannacea? corresponds with the hood of the Marantacese and not 
with the labellum of the Ziugiberaceaj. 

Order 4. Marantaceae. The flower is asymmetrical. Only 
1 or 2 of the 3 stamens in the outer whorl are present as stami- 
nodes ; in the inner whorl 2 are petaloid and of the sixth stamen 

FIG. 317. Calathea zebrina. 

one-half is developed as a staminode and the other half bears 
a bilocnlar anther. One ovule only in each loculus. The style 
is strongly curved and at first enclosed in one of the staminodes 
(hood) of the inner whorl ; later on it springs elastically forward 
towards the other staminode (inner-wing) of the same whorl. 
The stigma is very oblique or 2-lipped. Two of the three loculi 
of the ovary, in some (Maranta, Thalia) become small and empty. 
Embryo curved. Leaves in two rows, with sheath, stalk, and blade 
(Fig. 317) ; at the base of the last is a characteristic swelling 
(arf iculu>i) . Phrynium, Calathea, Stromunthe, Ctenanthe, Saranthe, etc. 
About 150 species ; tropical, especially America. The starch of the rhizome 
of Maranta arundinacea is OFFICINAL, " West Indian Arrowroot." 



Family 7. Gynandrse. 

The flowers are hermaphrodite and constructed on the ordinary 
3-merous, pentacyclic type with petaloid, epigynous, strongly zygo- 
morphic perianth, and generally one-stamened by the suppression 
of the other 5 stamens. The family has derived its name from 
the fact that the stamen is united with the style into a " stylar 
column " (except Burmanniacece,} . All are herbs; many grow as 
epiphytes on other plants. 

This family and the Scitamineas occupy correspondingly high positions 
among the Monocotyledons ; these two families may therefore be placed close 
together, although one cannot be derived from the other. The first of the two 
orders is very small, but the second is very rich in species. The Apostasies 
are best classed with the Orchidaceas and have no independent place. 

Order 1. Burmanniaceae. This order forms a transitional 
link between the Gynandra? and the epigynous Liliiflorse (Ama- 
ryllidacece), in having 1 a 6-leaved perianth, and 6-3 stamens ; but- 
some have a labiate perianth (the median perianth-leaf of the outer 
whorl being very large). The ovary is most frequently unilocular 
with three parietal placenta? ; but in some it is 3-locular with axile 
placentation. Capsule. Seeds oo, small, with endosperm. The 
relationship to the Orchidacere is shown especially in the very 
imperfectly developed embryo and in the ovary. Small, tropical 
herbs (59 species) ; some are saprophytes. 

Order 2. Orchidaceae. The epigynous, petaloid peria.nth is 

strongly zygomorphic in 
having the posterior leaf 
of the interior whorl, the 
lip (labellum), diffei-ing 
from all the other leaves 
in form, size, and colour 
(except Apostasies} ; the 
position of the labellum is 
very frequently reversed, 
being turned fonoards and 
downu-anls by the twisting 
of the ovary (Fig. 318 A). 
Only 1 of the stamens 
the anterior of the external 
whorl is developed and 
bears an anther (by the 
twisting of the ovary it is 


FIG. 318. A Diagram of an Orchid-flower. B, 
Cephalanthera. Stylar-column : a antlier; ,< stigma ; 
at the foot are seen scars indicating the position of 
the parts which have been removed. 

GtNANDRS. 329 

turned posteriorly and upwards) ; the others are entirely wanting 
(indicated by * in Fig. 318 A) or present as staminodes (Fig. 
318 A,<rcr) (except Apostasiece,Cypripedilex}; the filaments are united 
with the style to form a column (Fig. 318 5), the stylar-column l 
(gynostemium), and the anther (a) is thus placed on its apex and 
exactly behind or over the stigma (s). The anther is 4-locular ; 
the pollen-grains do not separate (except Apostasieue, Cypripedilece) 
but remain united either in tetrads or in masses, which correspond 
to a pollen-mother-cell (Fig. 320 C, D, E) or the pollen-grains, 
formed in each of the two anther-halves, remain united and form 
one or a few wax-like masses (pollen-masses, pollinia). The 3 
carpels form a unilocular ovary with 3 parietal, deeply bifid placenta? 
(except Apostasies, Selenipedilum). Only the two lateral carpels 
are prolonged and developed into the stigma (Fig. 318 7>, s), 
while the one lying in the median line, which is situated just 
within the anther (Fig. 318 A), becomes either rudimentary or 
developed into the " rustellnm '' (" a small beak "), on which the 
sticky bodies (glandule) arise ; by aid of these the heavy, connected 
pollen-masses may be glued to the insects which visit the flower, 
and pollination is thus secured (in Apostasiece and Cypripedilew the 
3 carpels each contribute to the formation of the stigma). The 
fruit is a capsule which most often dehisces by 6 valves, 3 of which 
are broader and bear the placenta?, and 3 alternating with them 
are narrower and barren (except Vanilla}. The very numerous 
and exceedingly small seeds have no endosperm, and have a some- 
what spherical embryo without any trace of external organs. The 
testa is membranous and loose. 

The Orchids are all perennial herbs with diverse habits and 
varying morphological structure (see the genera) ; the leaves are 
scattered, of the usual Liliaceous form, and the inflorescences in 
all cases are racemes or spikes (sometimes branched), with subtend- 
ing bracts, but without bracteoles. 

The forms which are the least modified are described first. 

I. APOSTASIES. The perianth-leaves are almost alike and free. 
The column is straight, with 3 equally-developed stigmas. Neu- 
wiedia has 3 perfect stamens (1 median of the outer whorl, and 
2 lateral of the inner whorl) ; Apostasia has only 2 perfect (inner 
lateral) and one barren (the median of the outer whorl), which how- 

1 According to Pfitzer, the column is the prolongation of the floral axis 
beyond the insertion of the perianth, and is not formed by the coalescence of 
sporophylls (filament and style). 



ever may be entirely wanting. The 3 posterior stamens are entirely 
suppressed. The pollen is powdery. The ovary is 3-locular with 
axile placenta. 7 species (Tropical East India, Australia). 

2. CYPRIPEDILEJ:. 1 The flower is strongly zygomorphic with a 
large boat-shaped labellum. There are two perfect stamens be- 
longing to the inner whorl, and the median anterior (later on the 
posterior) stamen of the outer whorl is transformed into a large, 
barren, shield-shaped body (Fig. 319). Selenipedilum has a 3-locular 
ovary, but Cypripedilum (Ladies'-slipper) has a unilocular ovary 
with 3 parietal placentae the typical structure for the Orchids. 

FIG. 319. Cijpripedilum calccolns : 1 front view of the flower ; 2 lateral view, after the 
removal of all the perianth-leaves with the exception of the labellum, which has been 
divided longitudinally ; 3 the slylar-column ; oo ovary ; s-s exterior, p interior perianth ; 
p' the labellum ; a the two fertile stameus ; a' the staminode ; st the stigma ; i entrance for 
the insects ; ex exit. 

The pollen-grains are separate (not in tetrads) and all the 3 lobes 
of the stigma are constructed to receive them. This group is there- 
fore, next to the Apostasieas, the least modified among the Orchids ; 
in all the following groups, one of the lobes of the stigma is differ- 
ently developed from the others, and there is only one stamen. 
Terrestial Orchids The pollination of C. calceolus is effected by the 
forcible entrance of insects into the boat-shaped labellum (Fig. 319 p') at /, aad 
their escape at ex (in 2) where the anthers are situated ; in this way the 
stigmas will first be touched and then the anthers. The pollen-grains are sur- 
rounded by a sticky mass in order that they may adhere to the insects. 

1 Cypripedilum = Cypripe.dium. 



3. NEOTTIE^:. The majority are terrestrial Orchids with creep- 
ing, sympodial rhizomes ; the blades of the leaves are not de- 
tached from the stem at joints, and have convolute vernation. 
The anthers do not drop off, but persist in the withered con- 
dition; their apex is brought in contact with the rostellum (acro- 
tonous Orchids). The pollen-grains are united in tetrads, which, 
however, often hang loosely together in pollinia, attached to a 
sticky part of the rostellum ("adhesive disc"), so that they 
adhere to the insects, and are by them transferred to the stigmas. 
Spiranthes. Listera ; Neottia. N. nidus-avis (Bird's-nest) is brown (it Las 
little chlorophyll) in colour, has no foliage-leaves, and lives mainly as a sapro- 
phyte ; the rhizome is studded with uubranched, fleshy roots which may form 
buds at their extremities. Vanilla climbs by aerial roots. The fruit 

FIG. 320. A Flower of Orchis maculate (front view) : a stamen ; b the cup ; 11 the 
stigmas; x staminodes; sp the spur; sji the entrance to it; srn-sl-sl exterior perianth- 
leaves ; )>m the labellum, and pl-pl the other 2 interior perianth-leaves. B-E Orchismascula : 
B lateral view of the column ; C a polliuium with massulse (p), caudicle (c) and adhesive 
disc (d) ; D caudicles with the cup (r), front view; the latter is depressed so that the 
adhesive disc is seen lying inside it ; E a pollimum, more highly magnified ; some massulfe 
are removed. F Oj'hri/s aranifera : rostellum and the base of the anther-loculus ; an 
adhesive disc is seen on the right. 

is fleshy and hardly opens, or does so irregularly. Epipactis, 
Cephalanthera. Epipogon and Limodorum are saprophytes without 

4. OPHRYDE^. Anthers 2-locular, not falling off, on a very 
short column. The anther is united at its base with the rostellum 
(basitonous Orchids, Fig. 320 A, J3), while in all other Orchids it is 
connected at the apex (acrotonous Orchids). The pollen-grains in 
each loculus are united into small "masses" (massulie), each of 


which corresponds to a polleu-mother-cell in the anther, and 
which hang together by elastic threads (Fig. 320 C, E). Each 
polliniuni is attached at the base by a stalk (caudicle) to an 
adhesive disc, formed by the modified stigma (rostellum), and is 
easily liberated from it (Fig. 320 C, D, F). The pollinium, which 
is formed in an anther-loculus, together with its caudicle and 
adhesive disc, is termed " pollinarium " (Fig. 320 C). All 
Ophiydete are terrestrial with tuberous roots, two of which are 
present in the flowering period, an older one (from the preceding 
year) containing the nourishment for the flowering-shoot of the 
year, and a young one which is intended to contain the reserve 
material for the following year. Inflorescence terminal. 

Orchis. The lip has a spur; each of the club-like polliuia is 
attached to its own adhesive disc, the discs being enclosed in a 
common pouch formed by the rostellum (Fig. 320 (7, -D). Tubers 
ovate, undivided: O. morio, mascula ; tubers palmate : 0. incarnata, macitlata, 
majalis. Ophrys; no spur, the two adhesive discs are each enclosed in 
a separate pouch (Fig. 320 -F). Anacamptis and Serapias have one 
adhesive disc. Habenaria, Gryvtnadenia, Platanthera, Herminium, 
Nigritetta,C(Kloylossiim,etc.,ha,ve naked adhesive discs (no rostellum). 

5. EPIDENDRE.E. Acrotonous Orchids with deciduous anthers (except 
Mida.ris) ; 2-8 wax-like pollinia, with or without caudicles ; generally no 
adhesive discs. Malaxis (the flower is twisted through a complete circle, causing 
the labellum to be turned upwards), Sturmia and Corallorhiza ' (Coral-root) ; 
the latter has a creeping, coral-like rhizome witnout roots, and is destitute of 
chlorophyll except in the ovary. The other two somesvhat resemble the 
tropical Orchids in having the lower interuodes of the axis of the inflorescence 
tuberous. Liparis; Calypso. Most of the genera are tropical epiphytes and 
many have aerial, green tubers formed from one or more steru-internodes ; 
Dendrobium, Eria, Pliains, bletia, Epidetidrum, Cattleya, Ltdia, Pleurothallis, 
Jiestrepia, Masdevallia, Bitlbopliyllum, etc. 

6. VANDE.E. These resemble the preceding but have only 2 wax-like pollinia 
in each anther, which are attached by a caudicle to the adhesive disc <>f the 
rostellum. Nearly all are tropical epiphyte?. Stanhopea, Catasetuin,Maxillaria, 
Oncidiiim 1'andn, Polystachya, etc. 

6,000 (10,000 ?) species. The majority live in the Tropics and occur, es- 
pecially, as epiphytes on trees or in the crevices of rocks, to which they are 
attached by aerial roots. These aerial roots, like those of Aracea?, are covered 
by several layers of spirally-thickened cells (tracheides) which contain air 
and form the velamen an apparatus to absorb moisture from the air. The 
roots have a white appearance when the cells are filled with air, which 
changes to a greenish hue when they are tilled with water, the chlorophyll 
then shining through. They generally have horizontal rhizomes ; the 

1 Corallorhiza = Coralliorrhiza. 



ascending shoots, which bear the foliage-leaves, may vary, but they very often 
swell and assume the form of a tuber, which persists for several years fresh 
and green after the leaves have fallen off (Fig. 321). Vanilla is an exception 
(see above). Our Orchids are all terrestrial (or marsh-plants) ; the largest 
number of species is found in calcareous soils. 

POLLINATION takes place principally by means of insects, but self-pollina- 
tion occurs in some. The lip serves as a landing-stage for the insect visitors, 
which, on sucking the honey, cause the adhesive discs, with the polliuia 
attached to them, to adhere, to their bodies (generally to the probosces) and so 
carry tbem away to other flowers. In some species parts of the flower are 
sensitive or irritable, which has some connection with the pollination. 
Without doubt there are a great many biological differences which are closely 
connected with the infinite multiplicity of forms ; Darwin (1862) has already 

FiO. 321. Chysis bractescens. 

shown an enormous variety, never even dreamt of before, in the European 
species. The genus Catasetum has $ - ? - and -plants with flowers of such 
different appearances that they have been classed in various genera (Myanthus, 
Monacanthus). Platanth?ra is pollinated by hawk-motbs ; Ophrys, by flies; 
Epipactis latifolin, by wasps ; Orchis, by bees, especially humble-bees, etc. 

The DISTRIBUTION OF SEEDS is effected by the wind, the seeds being so ex- 
ceedingly small and light. Many species moreover have peculiar, elater-like, 
fine, hygroscopic hairs in the ovary, which eject the seeds in a manner similar 
to the elaters of the Liverworts. 

The USES are few, mostly as ornamental plants in conservatories. The 
tubers of several Orchis-species are OFFICINAL ; they contain starch and mu- 
cilage and are used as " salep " The fruits of Vanilla pi ani folia are used as 
condiments and differ from other Ort'/t/'d-fruits in being rather fleshy and 
in dehiscing irregularly ; the seeds are very small, shining and black. 


Class II. Dicotyledones. 

In this class THE EMBRYO lias 2 seed-leaves, a rule from which 
there are few exceptions (e.g. Ficaria, Cyclamen, Pinguicula, 
certain species of Corydalis, with only 1 ; and a few, mostly para- 
sitic forms, e.g. Monotropa, Orobanclie, Pyrola, entirely without 
cotyledons). On germination the cotyledons nearly always raise 
themselves above the ground as green, assimilating leaves and are 
then termed aerial or epigean, in contradistinction to the under- 
ground or hypogean which are always buried. The structure of 
the seed varies (endospermous or exendospermous) ; the embryo 
may be straight or curved. In many instances the primary root 
grows as a vigorous tap-root, with weaker branches arising acro- 
petally (in annuals, biennials, many perennials, especially woody 
plants) ; but in a large number of herbaceous perennials, which 
have rhizomes, the root behaves very much as in the Monocoty- 
ledons. The roots generally increase in thickness by means of a 

THE STEM, when seen in transverse section, has its vascular 
bundles arranged in a ring; in reality, however, they form a 
kind of cylindrical network in the stem ; the bundles are open, 
and thickening takes place by means of a cambium ; annual rings 
are formed in the perennial stems. There is a rich and very varied 
form of branching. The two first leaves of a shoot (fore-leaves) 
are placed nearly always to the right and to the left ; the same 
arrangement is found in the two first leaves developed on the 
flower-stalk, and these are, as a rule, the only two ; they are found 
below the calyx and are usually termed the " bracteoles." It has 
become customary to indicate the bracteoles by the letters a and /3, 
according to their sequence of growth, and in that sense these 
letters will be employed in the following diagrams. 

THE ARRANGEMENT OF THE LEAVES varies very much ; there is 
also a great variety of shapes in the leaves and their venation, 
but the linear leaves, with parallel venation, so frequent in the 
Monocotyledons, are seldom met with, as also the large sheaths 
(though the sheath is well developed in the Umbelliferous plants); 
stipules occur much more frequently. 

THE FLOWER is most commonly cyclic, but acyclic or hemicyclic 
forms also occur. The type which may be taken as a basis 
consists in the majority of instances, as in the Monocotyledons, of 
5 whorls, of which the 4 outer ones (calyx, corolla, and the 2 


whorls of stamens) are most frequently 4 or 5 in number and 
placed in regular alternation, whilst the innermost one (the car- 
pels) has generally fewer members, probably on account of 
space (Figs. 360, 361, 421, 429, 487, etc.). Trimerous (Figs. 384, 
387, etc.) flowers, or those in which the members of the flower are 
in threes or a multiple of three, also occur, as well as dimerous 
flowers ; other numbers are rare. It is of the greatest importance 
in connection with the relative position of the members of the 
flower to the axis and bract (orientation), whether the bracteoles 
are typically present (even, thougfh they may not be developed), or 
are typically absent. If there are 2 bracteoles present, then their 
position in a pentamerous flower is often as follows: the first sepal 
turns obliquely forward, the second is posterior and median, the 
third obliquely forward, the fourth and fifth obliquely backward ; 
quincnncial aestivation is often found in these buds (Figs. 360, 429, 
471, 475, 584). The first and third leaves, in the following chapters, 
are most frequently alluded to as the " anterior," the fourth and 
fifth as the " lateral " leaves. The reversed arrangement, with the 
median sepal in the front, occurs for instance in Papilionacece 
(Fig. 511), Lobeliacece (Fig. 594), Rhodoracece. If any bracteoles are 
present below a tetramerous flower, the relation is generally that 
2 sepals (the first ones) stand in the median plane, the two next 
ones transversely (Fig. 393), and the corolla then adopts a diagonal 
position (Fig. 397) ; but a diagonal position of the calyx generally 
shows that the flower is not, strictly speaking, tetramerous, as in 
Plant ago (Fig. 567), Veronica (Fig. 559 C) and others. 

If the bracteoles are not typically present, then the position of 
the sepals is changed accordingly, and the two outer sepals en- 
deavour to assume the position which the bracteoles would 
otherwise have occupied, e.g. in Primula (Fig. 547). Other 
positions are also found when the number of bracteoles is more or 
less than two. 

The leaves which follow the sepals occupy definite positions 
with regard to them, which we may consider later. An arrange- 
ment must, however, be mentioned here ; when the flower is 
" diplostemonous," that is, has two whorls of stamens (thus, Su, Pn, 
An + n), these may be arranged in two ways. Either the first- 
formed whorl of stamens, which are termed the "calyx-stamens," 
stands directly in front of the sepals (that is " episepalous "), 
and is the outermost whorl, and in this case a regular alternation 
takes place between sepals, petals and the two whorls of stamens, 


which is also continued into the carpels if their number is the 
same as that of the other whorls : the carpels are then placed 
opposite the sepals (Fig. 278) and the flower is isomerous and Grn 
should be added to the formula above. Or, the calyx-stamens 
form the innermost whorl, and the corolla-stamens, which are sub- 
sequently formed (" epipetalous " stamens), stand outside these 
(Figs. 360, 429) ; if the number of carpels is the same as that of 
the preceding whorls, they are often placed right in front of the 
petals and the corolla-stamens. The first-mentioned arrangement 
is termed Diplostemonous, and the second Obdiplostemonous. Both 
arrangements may be found in one and the same order, e.g. Caryophyllacese. 
The size and relation of the members of the flowers, and also the contact with 
other members in the early stages of their development, play an important 
part in determining the arrangement. 

The great number of structural arrangements found in this 
enormously large class, may, as is the case in the Monocotyledons, 
be further varied by suppression and division of certain leaves 
(especially the stamens). Instances of this will occur in the 
following (Figs. 559, 568. 426, 441, 445, etc.). 

The Dicotyledons were formerly divided into 3 sub-classes : 
Apetala? (those without corolla), Sympetalaa or Gamopetala? (those 
with the petals united), and Choripetalaa or Polypetalae (the petals 
not united). This division has now been abandoned because it has 
been proved that the Apetalas were merely reduced or incomplete 
forms of the Choripetalse, and they have therefore been distributed 
among the various families of the latter sub-class. 

With regard to the Sym petals (or Gramopetalae) it may be 
stated that they form to a very great extent a closely connected and 
natural group, having in common not only the character that the 
corolla is gamopetalous and the stamens united with it (this being 
also found in the Cboripetalas), but also a great many others (such 
as persistent calyx, cyclic flowers with the formula S5, P5, A5 and 
as a rule Gr2, the two carpels being united to form the ovary ; seeds 
Avith a thick integument and a very small nucellus). They are 
therefore considered as an independent sub-class, and must be 
placed at the close of the system of classification as the forms 
which presumably have arisen the latest. In the future systems 
of classification this arrangement will very probably be changed, 
and the first families of the Sympetalee, the Bicornes and others 
will for instance be to a certain extent united with the families or 
orders of the Choripetalse. The Sympetala? may certainly be con- 



sidered as the youngest types, the strongly pronounced metamor- 
phosis supporting this theory, as also the formation of the integu- 
ment of the ovule, the one thick integument being undoubtedly 
derived from the coalescence of two a holochlamydeous ovule, etc. 

The Apetalte and Choripetalae are united into one sub-class. The 
leaves of the perianth in this case are, as a rule, free from each 
other, the structure of the flowers presents many differences, and 
the ovules have as a rule 2 integuments and a large nucellus. Con- 
siderable uncertainty still prevails regarding the arrangement and 
the relationship of the individual families of the Choripetalse, and 
some of the following families are hardly quite natural ; but the 
best arrangement arrived at so far has been adopted here. 

At the end of the book a review of the orders of the Dicoty- 
ledons will be found. 

Sub-Class 1. Choripetalae. Petals free. 
Family 1. Saliciflorae. 

Trees and shrubs, which, in the structure of the vegetative shoot 
and the catkin-like inflorescences, resemble the Querciflora?, bat 
the structure of the flower differs so much from them, that the 

FIG. 322. Male and female catkins of Sate caprea. 

only order brought under this heading Salicacece well deserves 
to be separated and to form a family of its own, the nearest 
relatives of which are still doubtful. As Juglandaceae and Myricacea? 
also deserve to be placed in a special family, the name Amentacea (Catkin- 
bearers), hitherto applied to all of these plants, cannot be retained as the name 
of a family. 

There is only one order. 

w. B. z 



Order. Salicaceae (Willows). Trees with simple, scattered, 
stipulate leaves. Dicecioiis. The flowers are arranged in simple inflor- 
escences (spikes or racemes) which are termed catkins, and which 
fall off as a whole after flowering ( <$ ) or after the ripening of the 
fruit (?) (Fig. 322). The perianth is very imperfect 1 or wanting, 
particularly in Salix (Fig. 323 o) ; the <$ -flower with 2 several 
stamens and without any trace of a carpel (a, fr, c) : the ? -flower has 
a free hicarpellate ovary, unilocnlar, and formed from 2 lateral car- 
pels with 2 parietal (median) placenta; and generally oo ovules; the 
style divides into two stigmas (d, e, f). The fruit is a two-valved 
capsule and the very small seeds bear a tuft of hairs at the base. 
Endosperm absent. The catkins are situated on dwarf-branches, which in 
some species often develop before the leaves arid bear at their base only 
scale-leaves ; in others foliage-leaves are borne beneatli the catkins. The vege- 

Fie. 323. Salix : male flowers of S. pentandra (a), S. aurlta (1), S. nibra (o); female 
flo\vers of S. aurita (d), S. ingricans (e), S. inollissima (f). 

tative bud commences with 2 bud-scales which are united on the anterior side 
into a scale. The capsule opens by the dorsal suture. The seed-hairs spring 
from the funicle. 

Salix (Willow) has short-stalked, most frequently lanceolate 
leaves and erect catkins with undivided bracts (Fig. 322). The 
flowers are naked ; 1 (o in a-f) or 2 yellowish glands situated in 
the median line. In the -flower generally two stamens, situated 
laterally like the carpels in the ? -flower. Various forms are seen in 
Fig. 323. The terminal bud of the branches often aborts regularly, the upper- 
most lateral bud taking its place. 

Populus (Aspen, Poplar) has long-stalked, more or less round or 
cordate leaves with drawn-out apex ; catkin pendulous ; lobed 

1 This is Eichler's view. According to Drude the perianth is absent ; at the 
base of the bracts, a nectary or cup-like disc. Prantl holds the same view. 
According to Pax the perianth is absent, but there is a disc cup-like, or reduced 
to a single toothed scale. 


bracts ; perianth cup-like with oblique edge ; stamens usually 
numerous ; stigmas often divided. P. tremuhi (Aspen) has received its 
name from the tremor of the leaves : cf. " to shake like an aspen leaf." 

POLLINATION. The Poplars are wind-pollinated. The Willows have sticky 
pollen and are pollinated by insects. The catkins of the Willows, especially the 
S , are more conspicuous, from the numerous, closely-packed, yellow flowers, 
rich in honey and pollen. The catkins often appear before the foliage and so are 
much more easily seen, whilst at this time of the year the number of competing 
honey-flowers is smaller, and the insect visits consequently more numerous. 
On many catkins of the Willow the flowers open earliest on the side which is 
turned towards the sun and in descending order, i.e. the upper flowers develop 
before the lower ones. Hybrids frequently appear. 

There are about 180 species existing in the northern, cold and temperate 
latitudes. Some in the Polar regions are scarcely more than an inch in height, 
and have a creeping rhizome (Salix herbacea, polaris, reticulata). Fossil forms 
are found in the Tertiary and perhaps also in the Upper Cretaceous. 

USES. Principally for ornamental trees, as they grow very quickly and are 
easily propagated by cuttings, S. babylonica, Weeping Willow; S.purpiirea; 
Poptdtis alba, Silver Poplar ; P. pyramidalis, Pyramid Poplar a form of 
P. nigra ; P. momlifera, Canadian Poplar. The wood is very poor and little 
used ; the branches of many Willows are cultivated for basket-making, etc. The 
wood of the Aspen is used for matches. The bark contains tannin and, in many 
Willows, a very bitter extract, Salicin (S. pentandra, fretgilis). Salicylic acid 
(officinal) is obtained from Sa.lix. Balsam is extracted from the buds of many 
Poplars, especially when the leaves are shooting. 

Family 2. Casuariniflorae. 

Trees with verticillate, scale-like leaves forming sheaths at 
the nodes. Monoecious. Flowers unisexual. -flowers in cat- 
kins ; ? in short spikes. Pollen-tube entering the ovule at the 
chalaza, and not through the micropyle. Ovary 1-seeded, 
unilocular. Carpels uniting into a multiple fruit. Only one 

Order. Casuarinaceae. Trees (30 species), from Australia 
and certain parts of S.E. Asia, with peculiar, equisetum-like 
appearance. The leaves are verticillate, scale-like and united into 
sheaths. The internodes are furrowed. Branching verticillate. 
The unisexual flowers are situated in catkins or short spikes. The 
c? -flower has a central stamen, surrounded by 2 median, scale- 
like perianth-leaves and 2 lateral bracteoles. The ? -flower has a 
1-chambered ovary (2 ascending, orthotropous ovules), no perianth, 
but 2 large, lateral bracteoles which finally become woody and 
form two valves, between which the nut-like fruit is situated. 
The multiple-fruits therefore resemble small cones. Casuarina 
equisetifolia, cultivated, gives " iron-wood." 


[The Casuarinas differ from the ordinary Dicotyledons in many important 
respects which may be briefly summarised thus : The bicarpellate $ -flower has 
a well-pronounced stylar-cylinder terminated by two stigmas, but the cavity of the 
ovary closes very soon after its formation, and in it are developed two parietal 
ovules ; these are united by a bridge of cellulose to the styJar-cylinder or summit 
of the ovary, and hence the ovules are connected with the walls of the ovary by 
the bridge (above), as well as by the funicle (below). The archespore is developed 
from the hypodermal cells at the summit of the nucellus, two primordial 
mother-cells are first formed and from these by tangential divisions a central 
cylindrical mass of cells (sporogenous-tissue) is produced which is surrounded 
by tapetal cells. The cells of the sporogenous tissue correspond to the mother- 
cells of the embryo-sac of other Augiosperms ; they divide transversely and 
from 16-20 macrospores are formed together with inactive cells which are not 
crushed together as in the case of other Phanerogams. The sexual apparatus 
is developed from a single cell, but the number of cells compo-iug this appar- 
atus is subject to variation, the oosphere being accompanied by one or two 
neighbouring cells which resemble canal-cells rather than syiiergidte. The 
sexual apparatus is found in the majority of the macrospores, but in most of 
these it remains as a number of .naked cells ; while in the fertile macrospores 
the cells are invested by walls of cellulose (usually only one fertile macrospore is 
found in each ovule). Antipodal cells are never developed. The macrospores 
elongate considerably towards the chalaza, into which some penetrate. The 
pollen-tube traverses the stylar cylinder and enters the ovules at the chalaza, 
its passage through the the tissue of the nucellus being assisted by the pro- 
longation of the macrospores. About the centre of the nucellus the pollen- 
tube is ruptured ; the apical portion which alone takes part in the fertilisation 
being firmly attached to the macrospore. Although the actual impregnation 
has not been observed, Treub considers that the endosperm begins to be 
formed before fertilisation.] 

Family 3. Querciflorae. 

Trees and shrubs with small, unisexual, monoecious flowers, 
having no perianth or a simple inconspicuous one. The <$ and ? 
flowers are very different and generally placed in separate inflor- 
escences. The ^-flowers are most often adnate to the bracts. The 
stamens are placed opposite the perianth-leaves, when they are pre- 
sent in equal numbers. The $ -flower is naked, or has a superior 
perianth. The ovary at the base is 2 or 3-(-6) Jocular with 1 or 
2 pendulous ovules in each loculus, only one of which is developed ; 
the fruit is a one-seeded nut ; endosperm absent ; embryo straight. 
The inflorescences, which are either compound and mixed (small 
dichasia in spikes) or simple, are here also termed catkins] but, 
strictly speaking, this term is applied to the <$ -inflor-escences only. 
In all Querciflorse the leaves are scattered (usually in 2 rows) 
simple, and penninerved, and with deciduous stipules. 



It is worthy of remark thcit in Betnlacea;, Conjlacea and Quercus the ovules, 
and to some extent the loculi of the ovary are not developed till after pollin- 
jition, so that the development of the pollen-tube proceeds very slowly. The 
smallness of the flowers, the absence of honey, the dryuess and lightness of tht 
pollen, the size of the stigma and the abundance of hairs found on many 
stigmas are all adaptations for wind-pollination. It is also an advantage that 
the flowers are generally pollinated before the foliage-leaves are developed, thug 
preventing the pollen being entangled by the leaves. 

The two orders Betulacece and Conjlaceie mentioned here are by other authors 
united into one order. [It is doubtful whether these two should be retained in 
the family Querciflorte, as recent researches (p. 273) have shown that they 
differ from the Cupulifera in many important points, and agree with tbo 
Casuarinas in the fact that the pollen-tube enters the ovule through thu 

Order 1. Betulaceae (Birches). Monoecious, with thick, 
cylindrical, compound $ and ? inflorescences (2- or 3-flowered 
dichasia in a spike with 
spirally- placed floral- 
leaves) (Figs. 324, 326, 
328). When the perianth 
in the ^ -flower is com- 
pletely developed, it is 
composed of 4 somewhat 
united leaves, which are 
placed opposite the 4 
stamens (Figs. 325, 326 A). 
The female flowers are 
naked ; the ovary is bi- 

locillar, with two Styles FlG 3M ._ 4llMW glut( : nosits . Branch of Alder with 

(J-(n) and <j>-(m) catkins: k bud ; It fruit-bearing 
catkin ("cone.") 


and one pendulous ovule in 
each loculus. The sub- 

tending floral-leaves unite with the bracteoles and form a 3-5-lobed 
cover-scale, which is not attached to the fruit (Figs. 325 D, 326 J3). 
Fruit a nut without cupule (see Corylacece and GupuUferce). In the 
bud the leaves are flat. The stipules are deciduous. On germination the 
cotyledons are raised above the ground. Terminal buds are only found on old 
Alder trees ; the Birch has svmpodial branches. 

Alnus (Alder) (Figs. 324-326). In the majority of species the 
(J- and $ -catkins are both developed in the year previous to their 
flowering, and pass the winter naked and bloom before the leaves 
expand. < -flower : 4 stamens. ? -flower : the 5-lobed cover- 
scales of the $ -catkin are woody and remain attached to the axis, 
so that the entire catkin when ripe resembles a small cone (Fig. 



FIG. 325. Alnus glutinosa: A dichasium of cJ -flowers seen 
from the front; B the same from inside ; C the same from the 
back ; D dichasium of ? -flowers with subtending-leaf and four 
bracteoles. The letters b, a, ft, ft', ft, are the same as in Fig. 
326 A. 

324 ?>). Each cover-scale supports two winged or wingless nuts. 
In the native species of Alder the buds are stalked (Fig. 324 k). The bud- 
scales are formed by the stipules of the lowest leaves. 

Betula (Birch). The <$ -catkins, in the native species, appear in 
autumn, the ? -catkins in the flowering year on leaf-bearing, short- 
lived shoots. (J -flowers : 2 stamens, divided (Fig. 328.4). The 3- 

lobed cover-scales 
(Fig. 327 a) of 
the ? -catkin are 
detached from 
the axis ; each 
cover-scale sup- 
ports 3 broadly- 
winged nuts (b). 
The stem has cork 
with annual rings. 
The young twigs and 
leaves have aromatic 
resin glands. 

In the axil of each 
cover-scale [b in the Figs ] is situated, in the $ -catkins (Figs. 326 A, 325 A-C) 
a 3-flowered dichasium, the flowers of which have a 4-partite perianth, the 
posterior perianth-segments being sometimes almost suppressed, and 4 stamens 
with undivided filaments. In the $ -catkin (Figs. 325 D, 326 C') a 2-flowered 

dichasium is found, the middle flower 
being suppressed (indicated by a star in 
C). In both instances the inflorescences 
have two bracteoles (a-/i) and the 
flowers borne in their axils have each 
one bracteole (/3'), the other one (a') be- 
ing suppressed and therefore in 326 A 
and C only represented by a dotted 
line ; these four bracteoles unite with 
the cover-scale (b) which supports the 
entire dichasium, to form the 5-lobed 
" cone-scale " (Fig. 326 B) which in the 
$ -catkin eventually becomes woody. 

A 3-flowered dichasium is situated in 
the axil of the cover-scale in both (?- 
and ? -catkins (Fig. 328 .4, B) ; only the 
central flower has bracteoles (a-j3) (the 
lateral flowers having no bracteoles), and these bracteoles unite, as in the Alder, 
with the supporting cover-scale (b), and form a three-lobed cone-scale (Fig. 327 a). 


FIG. 326. Alnus glutinosa: diagram 
of dichasia of $ (A) and ? (C) catkins; 
B a cone-scale. All the bracteoles in A 
and C are slightly pressed from their 
normal position. 


While the ? -flower exactly resembles that of the Alder, the reduction of the 
<? -flower, already described in the Alder, is carried further, so that often only 
the 2 median perianth-leaves are developed (Fig. 328 A) ; there are also only 
"2 stamens, these being deeply cleft, while the other 2 are suppressed. 

About 50 species ; N. Temp. Fossil-forms certainly occur in the Oligocene. 
During the Glacial period the Dwarf-birch (IS. nana) extended over Europe ; at 
the present time it is confined to the moors and mountains of N. Europe and 
N. America and Asia. Wind-pollinated. 

USES. Important forest trees. The bark contains tannic acid. The tar of the 
Birch is used in the preparation of Russia leather ; whilst its spring sap is very 
saccharine, and is used in some places for making a fermented drink. Its ex- 
ternal bark is used for roofing, for baskets, etc. 

PIG. 327. Betula tiernieosa : a cone-scale ; PIG. 328. Diagrams of dichasia in 

b fruit. the cj- (-4) and ?-(#) catkius of Birch. 

Order 2. Corylacese (Hazel-nuts). Monoecious. The - 
catkins are long and cylindrical ; the < -flowers are placed singly 
in the axil of the subtending-leaf (cover-scale) ; they are naked 
and formed of a number of divided stamens, which are partly 
united with the cover-scale, 4 in the Hazel, apparently 8 (Figs. 
330 A, 329 JB, G), more on the Hornbeam. The ? -flowers have 
a very small, superior perianth ; in the axil of each cover-scale a 
2-flowered dichasium (Fig. 329 D) is present, of which the 
central flower (* in Fig. 330 B) is suppressed. The gynoeceum is 
bicarpellary as in the Birches ; the ovary is bilocular, with two 
long styles (Fig. 329 D-F) the loculi have 1 (-2) ovules (Fig. 
330 -B). Each single ? -flower and fruit is surrounded by a leaf- 
like covering, the cupule (husk), which is formed of three floral-leaves 
(namely, the bract of a lateral flower, and its own bracteoles ; 
thus in Fig. 330 B, a, a', ft' form the cupule for the left-hand 
flower, and j3, a ; , /? the cupule for the right-hand). 

Gunjlus (Hazel-nut, Fig. 329). The long, cylindrical ^-catkins 
pass the winter naked, 2-3 together, on short branches. The very 



small ? -catkins are enclosed in buds, in which they pass the 
winter ; these buds are situated in the axils of the fallen foliage- 
leaves, and it is only by their larger size that they may be dis- 
tinguished from the ordinary foliage-buds. In spring the ? -catkins 
are easily recognised by their red, projecting stigmas (Fig. 329 A)- 
The cupule the "husk" is tubular, fringed, and envelopes the nut. 
The leaves are alternate and unsymmetrical, the external side being larger 
than the internal ; this is connected with the vernation, the blade being con- 

FIG. 329. Corylusavcllana : A branch at the time of flowering with <J-and $ -catkins; 
B cJ -flower with subtending-leaf (bract) and two bracteoles ; the same without the 
anthers; D view of interior of $-dichasium shortly after fertilisation; E young fruit 
with cupule ; F similar one with the cupule opened ; 6 mature $ -fruits ; H nut. 

duplicate in the bud ; the stipules are deciduous. The bud-scales are formed 
of stipules, the most internal having a leaf-blade attached to them which is 
suppressed in the external ones. The cotyledons remain underground on germi- 

Carpinus (C. betnhts, Hornbeam). 
appear till the leaves are 


The - and ? -catkins do not 
The $ -catkin in this 


instance is also long and cylindrical. The cupule in C. betnlns is 
3-lobed, and to a slight extent only embraces the base of the 
ribbed nut (Fig. 331) ; each lobe corresponds to a floral-leaf. 
Whilst the carpels are placed medianly in Corylus, in Caiyinus, on the other 
hand, they are situated transversely, as in the case of the Betulacea;. The 
lamina of the leaf is not conduplicate in the bud, but flat, and folded only along 
the lateral veins, which are also indicated in the form of the fully-developed leaf ; 
otherwise the vegetative characters are essentially the same as in the Hazel. 
The cotyledons are aerial. Oxtrya resembles the Hornbeam, but the cupule 
completely envelopes the nut, as a sac open at the apex (Eur., N. Am., Japan). 

N. Am., Asia, and Europe ; '25 species. Fossil forms in the Oligoceue. Wind- 
pollinated. USES. As timber (Carpinus betulus) and firewood. The fruits of 
C. avellana (ordinary Hazel-nut), C. titbtdosa (Lambert's nut) and C. column 
(Turkish Filbert) are edible. 

PIG. 330. Diagrams of the <5 -flower (A) of Corylus FIG. 331. Nut of the Hornbeam 

and the dichasiurn of the ? -flowers (B). with cupule. 

Order 3. Cupuliferse. Monoecious. The inflorescences make 
their appearance with the leaves, arising in the axils of the leaves 
of the same year. A woody cupule furnished externally with scales 
or spines is common, and surrounds 1-several flowers (the cupnle 
in the Corylacese never encloses more than a single flower or fruit). 
The ^-flower has a united perianth, which is, however, 4-6 partite, 
and encloses an indefinite number of undivided stamens. The ? - 
flower has a superior, 6-merous perianth (3 + 3, compare Figs. 
332 D, 334) ; the gynoeceum is formed of 3 (or in Castanea 4-6) 
carpels with a corresponding number of stigmas (Figs. 332 I>, 
H 334, 335) ; and the ovary has at the base 3 (-6) loculi (Fig. 
333), each of which has 2 pendulous anatr-opous ovules ; the fruit 
is a one-seeded nut (Figs. 332 E, 336). 

The cupule of the Cupulifera?, according to the opinion of 
Eichler, is formed by united bracteoles (compare Fig. 333, where 



the four valves in the cupule of Castanea are considered as 
bracteoles of the lateral flowers of the dichasium) ; according to 
another view (see Prantl, in Bugler's Bot. Jahrb., viii., 1889), it is 
a ring-like axial outgrowth independent of the bracteoles of the 
flower, whose scales arid spines are floral-leaves. The cupule in 
the Oak onlj encloses the base of the fruit, but in the Eating- 
chestnut and Beech the fruit is completely enclosed, and con- 

FIG. 332. Cnslanea vesca : A branch with inflorescences; B d -flower; C young cupule 
with three 9 -flowers; D $ -flower; E the same in longitudinal section; F cupule with 
3 nuts (diminished) ; 6f, H nuts (G in longitudinal section to shove embryo). 

sequently the cupule must divide into a number of valves (gener- 
ally 4) to allow the fruit to escape. In the 3-flowered dichasia 
of Pasania, Sect. Eupasania (Trop. Ind.), each individual flower 
has its own cupule of the same structure and development as in 
Qnercus, and, moreover, each group of flowers has externally the 
typical six bracteoles. 

Castanea (Eating-chestnut, Fig. 332). The catkins are erect 



(J.), cylindrical, with the ? at the base and the <$ at the top, or 
some are entirely $ and composed of small dichasia. The cupule 
(C, F) is 4-valved, provided with spines, and entirely envelops the 
3 nuts ; it is already developed at the time of flowering 1 . $ -flowers 
are most frequently borne in 7-flowered dichasia, aud have a well developed 
perianth, most frequently consisting of 6 leaves in two whorls (Fig. 332 />), and 
a large number of stamens. $ -flowers are most frequently borne in 3-flowered 
dichasia (Figs. 332 C, 333) ; the letters in Fig. 333 indicate the older theory, 
according to which the 4 bracteoles (a'-/3'j of the two lateral flowers are thick 
aud united into a single 4-valved, woody cupule, which surrounds the 3 nuts, 
and is furnished externally with spines ; the spines are well developed hair- 
structures. 6 carpels in two whorls. The leaves in the vertical shoots have 
a divergence of , |, $ ; on the horizontal shoots they are alternate. The 
cotyledons remain underground on germination. 

Fagus (Beech). The -catkins are pendulous, capitate; the <$ - 
flowers have an obliquely bell -shaped, fringed perianth, with 6-20 

FIG. 333. Diagram of the cupule of Castanea. FIG. 334. Female flower of Fagus (mag.) 

stamens. ? -catkins erect, 2-flowered, borne singly in the axil of 
foliage-leaves of the same year; the ? -flower has a gyuceceum 
formed of 3 carpels, bearing an epigynous, 6-leaved perianth (Fig. 
334). In this genus the dichasium has only 2 flcnvers, the central 
one being suppressed. The cupule contains, therefore, only 2 tri- 
angular nuts ('' mast "). All the shoots have the leaves arranged in two 
rows; the rows are on the underside, being only about 90" distant from each 
other ; the buds on the other hand approach each other towards the upper side. 
Tbe bud-scales are stipules without laminae ; in vernation the laminae are folded 
along the lateral ribs, the upper lateral portion being the largest (as iu Horn- 
beam and Chestnut). The cotyledons are folded, and at germination are 
aerial, large, and reniform. 4 species (Europe, Japan, N. Am.) Notlwfugus 
(S. Atn., New Zealand, S. Austr.) 

Quercus (Oak, Fig. 335). Catkins simple, ^-catkins long, 
thin, pendulous, few-flowered. $ -catkins erect ; the cupule is cup- 
like, entire, and encloses only the base of the solitary nut (" acorn "). 
The $ -flower has a similar construction to that of the Chestnut. The 



? -catkin has not more than 5 flowers (single-flowered dichasia, in which only 
the central flower is developed). The scales on the cupules are no doubt leaf- 
structures in this case also. According to another theory, the scales are hair- 
structures ; they arise on the internal face of the young cupule apparently in 
descending, but really in ascending order. The rirn of the cupule gradually 
expands. In the <j> -flower (Fig. 335) the loculi of the gynoeceum, together with 
the ovules, are not developed until after pollination. The leaves in all cases 
have a divergence of f ; the lowermost leaves on the shoots are reduced to 
stipules which serve as the bud-scales (5 rows). The laminae are conduplicate, 
as in Corylus, and the external side is the broadest. The cotyledons are fleshy 
and remain underground. 200 species. Paaania (100 species). 

368 species, in temperate climates, especially in Europe and N. America. 
Authenticated forests have been found in the Oligocene. The Beech has one 
species, Far/us sylvatica, in Europe ; it is a most important forest tree (in 
Denmark the most important) and reaches its most northern limit near Alve- 
sund in Norway (60 N.L.), its northern boundary line passing from Alvesund 
in a zig-zag line through Ludwigsort, south of Konigsberg, in Prussia, to- 
wards the Crimea. According to Steenstrup and Vaupell, the Beech did not 


FIG. 335. Quercus : A 9 -flower in its cupule 
(mag.); B longitudinal section through A, 
showing cupule, perianth, and inferior ovary. 

FIG. 336. Fruit of Quercus. 

make its appearance in Denmark until a comparatively recent time, the Oak 
then being partially supplanted. Other species of Beech are found in N. 
America and Japan. Several species of Nothofagus occur in the South West 
of S. America, and in the colder regions of the southern hemisphere. The 
Oaks grow especially in temperate regions, e.g. in Western Asia, N. America, 
and the mountains of Mexico. Evergreen species are found in Tropical Asia, 
Himalaya, Japan and the Mediterranean region. In this country there is one 
species of Oak (Q. robur), of which there are three varieties (Q. pedunculata, 
intermedia, sessili flora). The Eating-chestnut is found in the South of Europe, 
but is cultivated in the midland and southern counties of England. USES. 
The wood of these trees is very useful as timber. The wood of Q. tinctoria 
has a yellow colouring matter (Quercitron-wood). The bark of the Oak contains 
a large quantity of tannic acid, and is used for tanning ; for this purpose 
also the cupules of Q. vallonea, (f/jilops, grteca, and others from the Eastern 
Mediterranean, are used under the name of " Yalloons." The Cork-oak 
(Q. suber ; S.W. Europe) is the most important tree from which cork is obtained, 



its bark being very largely developed and stripped for cork. Gall-nuts are found 
on many species ; those of Q. lusitanica, var. infectoria (Eastern Mediterranean) 
are officinal, and likewise the fruits (acorns) and the bark of Quercus peduiicu- 
lata and sessili flora. Oil is obtained from the Beech " mast." The nuts of the 
Chestnut tree are edible. 

Family 4. Juglandiflorse. 

This family resembles the Quercifloras in the catkin-like inflores- 
cences, the imperfect, unisexual flowers, the epigynous perianth 
and the woody shoots with scattered leaves, etc., though it is in 
other respects very dissimilar ; one point of difference is the presence 
of aromatic compounds, but a more imporant divergence is found 
in the structure of the gynoeceum, which is formed of two carpels 
with one loculus and has one basal, orthotropous and erect ovule, 

FIG. 337. Juglans regln : A <J -flower seen from below with bract (cover-scale) (6), 
bracteoles (a and /3), perianth-leaves (p) ; B the same from the front ; C lateral view of the 
same; D diagram of A ; E $ -flower with bract, the bracteoles are united with the ovary, 
their edge being visible as an indented line below the perianth ; F 2 $ -flowers at the end of 
a foliage-shoot; Gf fruit (without the fleshy covering,) in longitudinal section; H trans- 
verse section of the same. 

which, as in the Querciflorre, does not become developed until 
after pollination ; the fruit too is very different, being generally a 
drupe. Endosperm absent. 

Order 1. Juglandaceae (W^alnuts). Leaves sea tiered, impari- 
pinnate, rich in aromatic compounds. Stipules absent. Flowers uni- 
sexual. Monoecious. The catkins are lateral, generally on naked 
branches of the previous year, cylindrical, pendulous, many-flowered; 
the two bracteoles and the 2-4-leaved perianth of the <$ -flower unite 
with the subtending bract; the ^-fkwer- has indefinite stamens 
(6-20 in Juglans, Fig. 337 A-D) . The ? -catkins are terminal, gene- 
rally on branches of the same year, few-flowered (Fig. 337 F) ; the 



? -flowers have a superior, 4-leaved perianth, a bicarpellate gynre- 
ceum, two styles with stigmas on the internal surface. The ovary, 
bracteoles and bract all unite together (Fig. 337 E). The fruit is 
generally a green or black drupe, 1 whose flesh (outer soft portion) in 
Gary a and Juglans ruptures more or less irregularly, and frees the 
stone ("Walnut"). The stone in Juglans is divided internally by one true 
(Fig. 337 H) and by several false, low partition walls into several incomplete com- 
partments, so that the two large cotyledons become lobed and incised to fit like 
a cast into the irregularities of the inner surface of the stone ; the embryo is 
exeudospermous and covered with a thin testa. THE LEAF SCABS are large and 
cordate with 3 groups of vascular bundles. The PITH in Juglans and Pterocarya 
is divided into chambers. The stone ruptures, on germination, along the 
dorsal suture into 2 valves ; the cotyledons remain underground. In Juglans 
regia a long row of accessory buds is found on the lowest internode (epicotyl) 
above the axils of the cotyledons. Pollination by the wind. Both protogyuous 
and protaudrous examples of Juglans re<]ia occur. 33 species, mostly in tem- 
perate North America. USES. Walnuts are obtained from J. iiigra and regia ; 
Hickory from North American species of Canja. The oil-containing seeds of 
several species are edible. Pterocarya and others are cultivated as ornamental 

FIG. 338. Myrlca gale : a young fruit ; x the bracteoles with numerous glands ; I longi- 
tudinal section of fruit. 

Order 2. Myricaceae. To this order belong shrubs or trees 
which have penninerved, simple, at most lobed or pinuatind leaves, 
with or without stipules, and with yellow, aromatic, resin glands 
(Fig. 338 a). The flowers, situated in catkin-like spikes, are 
unisexual and naked, and supported by scale-like floral-leaves, 
cj -flower: 4-6 (-16) stamens with short filaments; ? : generally 
situated singly. The oynceceum has a short style with 2 long 
stigmas, and unites with the bracteoles, which form wing-like 
outgrowths on the ripe drupe as in Pterocarya in the Juglandaceee 
(Fig. 338). Cotyledons fleshy (Fig. 338 6). Myrica ; Comptonia. 

1 The fruit of the Walnut is thus a false fruit ; and the term drupe must 
therefore not be used in the same sense as in the Eosacea?. 


40 species ; Temperate. Myrica gale (Sweet-gale, Bog-myrtle) has been used 
in the preparation of beer (Sweet-willow beer) on account of its resinous essential 
oil. M. cerifera (N. America) and species from the Cape, M. quercifolia and 
others, form wax on the fruit which is used in the preparati >n of candles. 

Family 5. Urticiflorae. 

The flowers are regular, hi/pogynous, nearly always unisexual, 
small and insignificant, Avith single, green perianth of 4-5 leaves. 
Stamens 4-5, placed opposite the leaves of the perianth. Ovary 
formed of 1 or 2 carpels, most frequently tmilocular, with one 
ovule (Fig. 340). The fruit is a, nut, more rarely a drupe, with one 
seed, generally endospermous. The Nettles are the sole order in the 
family which has only one carpel (1 stigma) ; this turns the posterior side to 
the front (Fig. 340). The others have two carpels (2 stigmas) but the anterior 
only is fertile (Fig. 346) except in a few Ulmacese and Moraceas. 

The majority are trees or shrubs with petiolated leaves, stipu- 
late ; rough hairs are very frequently developed upon the leaves. 
The flowers are very often crowded together in the inflorescence, 
which is rarely catkin-like. Peculiar aggregations of fruits are 
found in some orders. Latex and tough bast, which is used techni- 
cally, are also frequently found. Cystoliths are found in the 
epidermis of many species of Ficus, Urtica, and others. Wind- or 
self-pollination is most common, as in the Querciflor* and Juglandiflorse. In 
the Urticacese, 3L>rus and some others, the stamens lie incurved in the bud, 
and when ripe straighten themselves suddenly and elastically, and thus small 
clouds of pollen-grains are ejected with considerable violence on to the stigmas, 
which are often provided with brush-like hairs (Fig. 341). The formation of 
honey does not take place. 

Order 1. Ulmaceae (Elms). Trees or shrubs without latex. 
Leaves simple, arranged in two rows (divergence i), oblique (the 
inner side, nearer the axis, being the larger), strongly penni- 
nerved, dentate, hispid; stipules deciduous. In opposition to the 
other Nettle-like plants the flowers are often with a united 
cup- or saucer-like, generally 4-(5)-6-divided perianth, and a 
corresponding or larger number of opposite erect stamens. The 
gynoeceum has two carpels (2 stigmas), generally one loculus with 
one pendulous, anatropous or amphitropous ovule, 1 seldom two 
loculi and 2 ovules. Fruit one-seeded (nut or drupe). Embryo 
without endosperm. 

A. ULMK.E. The fruit is a ivinged nut (Fig. 339), the embryo 
straight, without endosperm. Anthers extrorse. Ulmus (Elm). 

1 The pollen-tube in Ulmus does not enter the ovule through the micropyle. 



The flowers are situated in inflorescences which develop from 
the lower buds of the shoot of the preceding year. The lowermost 
bud-scales are empty, the uppermost support either solitary flowers, or small, 
dichasial or unipared scorpioid inflorescences. The terminal bud on tbe vege- 
tative shoot quickly falls off, and the 
upper lateral bud continues the growth 
sympodially. Flowering takes place 
before the leaf-buds open. The flowers 
are wind-pollinated and have no honey. 
Fossil species have been found in the 

20 species ; North Temp. (2 species 
in this country). Important as timber. 
The Cork-elm (U. sttberosa) has a rather 
thick cork, which, however, is of no 
technical use. The bast is used as 

B. CELTIDE.E. The fruit is a drupe, 
the embryo curved, with folded or rolled 
up cotyledons, with or without endo- 
sperm. The anthers are introrse. The 
flowers are borne on a shoot of the 
same year. Planera (N. America) ; 
Zelkora. About 114 species; especially 
N. Temp., Trop. 

FIG. 339. A Ulmus campestris, flower 
with exceptionally aborted g.ynoeceum; 
B, U. effnaa, flower with 8 stamens ; C, U. 
cuiiipe-ftn's, fruit opened in front to show 
the seed pendulous from the apex of the 
loculus ; one loculus is aborted. 

Order '2. Urticaceae 

(Nettles). The majority of spe- 
cies are herbs with simple, stipu- 
late leaves ; they have no latex ; 
stinging liairs abundant. The flowers (Fig. 340) are unisexual, gene- 
rally 2-meroQS and arranged in clusters, -which are united into 
catkin-like inflorescences. The perianth is composed very often of 
4 (2 + 2) free, or in the ? -flowers generally united, green leaves ; 
the 4 (2 + 2) stamens are opposite the perianth-leaves, the filaments 
are bent inwards in the bud and throw themselves elastically 
towards the outside. The gynoeceuni has one style and one stigma 
(capitate or brush-like, Fig. 341) ; the ovary is unilocular, with one 
orihotropous, erect ovule (all other orders of this family have inverted 
or curved ovules). Fruit, a nut or drupe. Endosperm present 
(in Urtica very little), oily. Embryo straight. The STINGING HAIRS 
are club-shaped, very turgid, and provided with a siliceous, brittle apex, 
which breaks off in an oblique direction and allows the poisonous cell-sap to 
be forced out. In many tropical Nettles this is so strong that it may produce 
partial paralysis. There is no rudiment of an ovary in the $ -flowers (Fig. 3-40 
A). The PERIANTH in the ? -flower differs from that of the $ in having the two 


internal leaves generally much larger and enveloping the fruit (340 B) ; it 
often happens that all the perianth-leaves are united to form a gainophyllous 
envelope. -flowers may occur among the others. THE INFLORESCENCES among 
our native species are dichasii, which become transformed into unilateral 
scorpioid cymes by the development of the bud of the 2nd bracteole. In 
Parietaria they are more pressed together, and the floral-leaves at the same 
time are also raised on their axillary shoots to just beneath the flower. As a 
rule, not only in this order but also in those related to it, a small vegetative 
branch is situated in the axil of the foliage-leaf, and this bears an inflorescence 
on each side at its base. 

Urtica (Nettle) has opposite leaves with distinct stipules and 
stinging hairs. The perianth-leaves of the ? -flower are free 
(Pig. 340). -Parietaria (Pellitory) has scattered leaves without 
large stipules, and stinging hairs are absent. The ? -perianth is 
4-toothed, flask- or bell-shaped. Pilea is a tropical genus with trimer- 
ous, zygomorphic ? -flowers, the posterior perianth-leaf being much larger than 

FIG. 310. Diagram of 3- and ? FIG. 311. Parietaria dijf'usa; hermaphrodite 

flowers of Uriica dioica. flower : a in the female, b at the commence- 

ment of the male stage ; the stigma lias fallen 
off, but the anthers have not yet dehisced. 

the two others, and more or less hood shaped. The flower of Forskohlea is the 
most reduced; the $ -flower has only one stamen, and the ? - as well as the $ - 
flowers have a one-sided, tongue like perianth (?). Pouzolzla. 

WIND-POLLINATED. The pollen is shot out of the anthers, when they spring 
forward, and is caught by long stigmatic hairs. Parietaria diffasa is protogy- 
nous (Fig. 341). 

500 species ; chiefly in the Tropics, although the few species which occur 
in Europe are represented by a much larger number of individuals. USFS. 
The bast of the native species Urtica dioica and urens, of U. canuabina 
(Siberia), etc.; of Boehmeria nicea "Ramie" and " China-grass" (from Sunda 
Is., China), and others, is used in the manufacture of muslin. 

Order 3. Moraceae, (Mulberries). Nearly all trees or 
shrubs, seldom herbs, generally with latex. The leaves are scat- 
tered, and not infrequently lobed. The flowers are unisexual 
(monoecious or dioecious) and arranged in catkin- or capitulum-like, 
compound inflorescences. Perianth-leaves 2-6, generally 4, with an 
equal number of stamens opposite to them, as in the Nettles. The 

w. B. A A 



ovary is 1- seldom 2-locular, and has 2 stigmas (it is thus formed 
from 2 carpels) seldom only one style with one stigma. One 
ovule in each loculus, more or less curved, and pendulous ; micro- 
pyle directed upwards. Fruit usually a drupe. The embryo is 
generally curved inside the fleshy endosperm, or it is exendospermous. 
A. MOKE.S. The filaments are incurved in the bud. Leaves 
folded in the bud -Morns (Mulberry) (Figs. 342-344). Monoecious. 
The inflorescences are catkin-like in appearance, but in reality 
composed of many small dichasia. The flowers are similar to those 
of the IVettle, but with 2 carpels : in the < with perianth 2 + 2, 
and stamens 2 + 2 (Fig. 342), in the ? , perianth 2 + 2, and 2 
carpels in regular alternation. The small drupes are enveloped 
by the perianth, ivhich eventually becomes fleshy, and as all the 
flowers on the axis very accurately fit together, the collection of 

FIG. 342. Morns alba 
<J flower (611). 

FIG. 343. Morns alba 
$ inflorescence. 

FIG. 34!. .Moi-us nigra 

fruits is formed, which we call a Mulberry (Fig. 344). The leaves 
are folded in the buds, and have small stipules. The following are 
allied to Morns : Madura, Bromsonetia (the Paper-mulberry tree) which has 
spheroid ? inflorescences (made up of dichasia), etc. 

Dorstenia presents an interesting transitional form to the Fig in its flat, 
open, and, in some instances, lobed inflorescence on which the $ and $ 
flowers are sunk in grooves. Indications of a somewhat similar structure are 
found in certain Nettles, the sympodial axes of the dichasia becoming flatly 
expanded. The fruits are 1-seeded, but, nevertheless, spring open and eject 
their seeds. 

B. AKTOCARPEJ:. Filaments straight in the bud ; foliage- 
leaves with convolute vernation. An interpetiolar leaf-sheath 
(ocrea) formed in the axil of each leaf by the connate stipules, 
covers the younger leaves as a hood. It falls off as the leaf 
expands, and leaves a ring-like scar on the stem. Ficns (the 
Fig). The inflorescence (the so-called syconus) has a pear- 



shaped, fleshy, but hollow axis, on the interior surface of which the 
flowers are situated (Fig. 345). It is a kind of capitulum, with 
a hollow receptacle, whose " involucral " leases close over the 
entrance to the interior ; it is not, however, a simple capitulum, 
but a coalescence of cymose inflorescences. The edible parts are 
the fleshy stem-portion and perianth-leaves. The <$ -flower 
has a 2-6 divided perianth, 1-2 (-6) stamens ; the ? -flower 
has an oblique ovary. The fruits are drupes, with thin flesh. 
Many species have aerial roots, and some live as epiphytes on trees. 
POLLINATION, in the edible Fig, is effected by a small Gall-wasp (Cynips 
psenes L.), which lays its eggs in the Fig, and hence carries the pollen away. 
Even in very ancient times it was 
customary to hang infected wild Figs 
on the branches of cultivated ones, so 
that the young Gall-wasps, as they 
emerged, could immediately effect 
the pollination (caprification). Ficus 
carica, and other species, have two kinds 
of ? -flowers, besides the $ -flowers. 
One kind has a short style and no stig- 
niatic hairs, and it is only in the ovaries 
of these that the wasps lay their eggs 
(gall-flowers) ; the other kind has a long 
style and well-developed stigmatic-hairs, 
but the wasps cannot reach their ovaries 
these are "seed-flowers." There are, 
moreover, two kinds of plants of Ficus 
carica ; $ -plants, which have only seed- 
flowers, and bear the edible Figs, and 
<? -plants (called " Caprificus "), which 
bear inedible fruits, and have $ -flowers 

at the upper part of the Fig, but gall- 

mu /-i -c FIG. 345. A Fier in longitudinal section. 

flowers at the base. [The Caprificus, at 

Naples, bears three crops of inedible Figs each year, viz. Mamme (April), Pro- 
fichi (June), Mammoni (August). The $ -flowers are produced especially in June, 
the first Figs being almost entirely $ , and the last having but few $ -flowers. 
Each crop produces a new generation of Fig-wasps. The female wasp enters 
the Figs on the Caprificus, and lays one egg in each flower, with the result that 
the flower developes into a kind of gall. The mother-wasp dies within the Fig. 
The male wasp is wingless ; it bites a small passage into the ovaries containing 
the female wasps, and impregnates them ; the female wasps theu escape from 
the Fig, those in the Proficlii carrying pollen away with them as they pass out. 
They then enter another Fig, lay their eggs, and die. The edible Fig-tree 
similarly has three crops in the year, Fiori di Jico, Pedajnuali, Gimaruoli. 
The wasps, entering these Figs, are unable to lay their eggs in the ovary, but, 
nevertheless, they effect cross-pollination on entering the Pedagnuoli, which' 
bear fertile seeds.] 


The flowers of Brosimum are the most reduced. The perianth is -wanting, 
and the $ -flower has only 1 stamen. Cecropia (Trumpet-tree), in S. Am., has 
its pith divided into chambers ; these are inhabited by ants, which feed upon 
small food-bodies formed on the swollen base of the petioles. The leaves are 
petiolated, often shield-like, fringed or lobed, and sometimes with white felted 
hairs. They serve as food for Bradypus (the Sloth). Sorocea ; Castilloa. 

About 300 species exclusively in the warmer climates. The white Mulberry 
(M. alba, from China, India, Mongolia) is cultivated for the sake of its leaves, 
which are the indispensable food for silkworms. The black Mulberry (M. nigra, 
W. Asia) is cultivated for its fruits, which are used for the officinal Mulberry 
juice. The ordinary Fig-tree (Picas carica) is from the Mediterranean. The 
fruit of the well-known Oriental Sycamore (/*'. sycomorus) is edible. The Bread- 
fruit tree (Artocarpus incisa) and the Jack (A. integrifolia) have their home 
in the South Sea Islands, and are cultivated in tropical countries. The 
Bread-fruit is morphologically the same as the Mulberry. It has a very large, 
spheroid inflorescence, whose floral-leaves and perianth become fleshy and 
united into one nutritious mass, together with the axis, which is also fleshy. 
The milky juice of the India-rubber tree (Fie us elastica, East Indie*, a com- 
mon house-plant), and of Castilloa elastica (Am.) is the raw material of India- 
rubber. The milky juice of Galactodendron utile. (Cow-tree, S. Am.) is sac- 
charine and nutritious, but in Antiaris toxicaria (the Upas-tree, of Java) it 
is a strong poison. The bast of the Paper-Mulberry tree (Dr. papyrifera, 
Eastern Asia; is used in Japan for paper. Shellac is obtained from a small, 
hemipterous insect (Coccus lacca), which lives upon Ficus laccifera and F. 
rcligiosa (the Bo-tree, sacred to Buddha), E. India. The wood of Madura 
aurantica (Am ) has a yellow colour, and is known as yellow Brazilian wood. 

Order 4. Cannabaceae. The plants which belong to this 
order are aromatic herbs, either annuals or perennials, without 
latex. Leaves palminerved, and more or less divided, hispid, and 
with free, persistent stipules. Flowers always dioecious; ^-flowers 
in panicles, formed of dichasia, passing over into uniparous scor- 

pioid cymes. They differ from the 
Nettles, particularly in the 5-leaved 
perianth of the ^-flower, the 5 sta- 
mens (Fig. 346-351) with filaments 
erect in the bud, and in the ? -flower 
by the small, entire, cup-like perianth, 
FIG. 346. Diagram of male and which surrounds the base of the ovary 

e flowers of the Hop and (pig. 346, p. 352). The ovary has 

Hemp : b the bract, p the perianth. . . - . 

The position of the embryo is tw o styles, or one divided into two, 
indicated. with two stigmas and a pendulous, 

curved ovule (Fig. 346 B, 352 B) ; the fruit is a nut ; the embyro 
is curved (Hemp, Fig. 353), or rolled (Hop, Fig. 349), ivithout 



Only 2 genera with 3 species (Asiatic), of which two are culti- 
vated. Humulus lupulus (Hop, Figs. 347-349) is a twining, 
perennial plant, twisting to the right, with opposite, palmilobed, 
rough leaves, and large, interpetiolar stipules. The ? -flowers 
are situated in closely-flowered, cone-like, compound inflores- 
cences, with ultimately large, thin, imbricate floral-leaves (Fig. 
348) which bear the yellow, glandular hairs, containing lupulin. 

FIG. 317-318. Humulus hipnius : 347, twining stem ; 348, branch with strobiles. 

This inflorescence is made up as follows : The most external floral-leaves are 
situated iu pairs, an! are the persistent stipules of a leaf, the blade of which 
has become suppressed, or in any case is rudimentary. Such a pair of stipules 
supports 4 ( - 2-6) flowers in a double uniparous cyme, whose central axis does 
not develope into a flower. The bracts of these flowers (bracteoles of the 
partial inflorescence) become, at maturity, very large, spathe-like, and, together 
with the stipules, produce a cone-like appearance. 

Cannabis sativa (Hemp, Figs. 350-353) is an East Indian herb, 
with palmilobed leaves, and differs from the Hop in being annual, 



erect, and in having its leaves opposite at the base and scattered 
above. The $ -inflorescence is not cone-like as in the Hop, but the 

flowers are similar in construction. The main 
difference is to be found in the axillary shoot, which was 
suppressed in the Hop, and is in the Hemp developed 
into a leaf-bearing shoot which on each side bears only 
one ? -flower, and in the fact that the bracts are not so 
strongly developed. 

The " Hops " (the female inflorescences) are used in 
brewing, and medicinally on account of the yellow 
glands which contain lupulin. The Indian variety of 
Cannabis sativa contains an abundance of glandular 
hairs and resin. The withered inflorescences are used 
in medicine and are officinal. The bast of the stems 
of the Hemp is also used and the fat oil of the seeds. In Oriental countries 
the entire plant is used in the preparation of an intoxicating drink (haschisch), 
the narcotic material being found in the glandular hairs. 

FIG. 319. Humulus 
lupulus: fruit in longi- 
tudinal section. 



FIGS. 350-35i. Cnwnnbis sativa: 350, -plant; 351, (J -flower; 352, $ -flower, entire and 
in longitudinal section; 353, fruit in longitudinal section. 

Family 6. Polygoniflorae. 

This family is on one side closely allied to the Urticacece by its 
solitary, basal, vertical, and straight ovule, and by the conical ocrea 
which envelopes the younger leaves in the bud, similar charac- 


ters being present in the Urticacefe. On the other side it is 
related to the Curvembryae. The flowers are small, often 
trimerous, regular and slightly perigynons (in Chloranthacea, if they 
properly belong to this family, and Houttuynia, more or less epigynous). 
Syncarps are present in some Piperacea?, but the fruit is generally 
a single fruit, one-seeded berry, nut or drupe. The leaves are 
generally scattered. 

Order 1. Polygonaceae. The majority are herbaceous plants 
with round, often jointed stems, scattered leaves and ocrea, that is 
a membranous, tubular, ligular or stipular structure inside the 
base of the leaf, which clasps the stem and axillary bud ; the 
edges of the lamina are rolled backwards in the bud. The flowers 
are regular, small, generally , slightly perigynous, with incon- 
spicuous, simple, green or white perianth of 5-6 free segments; 
stamens 5-9 (Fig. 354) sometimes 
arranged in two series ; gynoeceum 
2-3 carpels, ovary unilocular with 
fine basal, straight (orthotropous) 
ovule, 2-3 free styles. The fruit is a 
2-3-angular nut ; the embryo, with 
mealv endosperm, is straight or 
curved (Fig. 355 /f), often unsym- 

metrical. The inflorescences are com- 
pound, and generally branch from the 
axils of the bracteoles, so that the last 

partial-inflorescences become coiled, uui- FIG. 361.-J Diagram of IMeum ; B ot 
... r, i Rumex; C of Polygonunt faqopyrum; T> 

parous scorpioul cymes; in Polygonunt .., ' mu 

J of P. lapabhifoliwn. The ovules are in- 

the two bracteoles unite into a mem- d i catec i inside the ovaries ; bracts and 
branous tube ; in Rheum and Rumex there bracteoles are not shown, 
is only one bracteole. 

Rheum (Rhubarb, Fig. 354 A) has a 6-leaved, petaloid perianth 
(Pn 3 + 3) and 9 stamens (A3 3 + 3). The 3-winged nut is nut 
enclosed by the perianth. 

Rumex (Dock, Fig. 3545) has 6 stamens (A3 2 + 0); the perianth 
is 6-leaved (Pr3 + 3), green or red, and the triangular nut is en- 
veloped by the 3 interior perianth-leaves, which point upwards 
and continue to grow after flowering. These perianth-leaves often 
have warts on their outer surface. The following are monoecious: R. 
acetosa and R. acetosella. 

Polygonum (Knot-grass, Figs. 354 C, D; 355). The petaloid 
perianth is most frequently 5-merous (| spiral) ; 5-8 stamens. 
The nut is triangular (Fig. 354 C, 355), or lenticular (Fig. 354 D). 



There are two whorls of stamens, the external with introrse, and the internal 
with extrorse anthers. The gynoeceum is often bi-carpellate (Fig. 354 D). 

The flowers may be considered as coustmcted upon the monocotyledonous 
type. Pterostegia has a perfectly monocotyledonous flower with 5 trimerous 
whorls. Rheunt likewise, bat here the external staminal whorl is doubled (Fig. 
254 A). Oxyria has a dimerous Eftewm-flower (4-leaved perianth, 6 stamens, 
2 stigmas). Rumex has a Rheum-Rower with the suppression of the internal 
whorl of stamens (Fig. 354 Bj ; Ernex is a dimerous Rumex. Polt/gonum, to 
which Coccolobn, Miildenbeckia and others are related, differs from Rheinu 

FIG. 355. Polygonum fagopyrwn : A branch with flower and fruits (nat. size) ; B flower ; 
C the same in longitudinal section ; D anterior and posterior view of stamen ; E gynoeceum ; 
F fruit (mag.) ; G fruit in longitudinal section; H transverse section, showing the curved 
cotyledons embedded in the endosperm ; I the embryo. 

chiefly in having one of the leaves, which in the latter takes part in the forma- 
tion of the perianth, developed in this case into a bracteole (so that the perianth 
is reduced to five members), and several or all the stamens in the inner whorl 
become suppressed. The perianth in Coccnloba and Miihlenbeckia is more or 
less perigynous and becomes fleshy, enclosing the fiuit. Miildenbeckia 
platyclada has flat branches with rudimentary leaves ; sometimes branches 
with normal, arrow-shaped leaves are found. Atraphaxi*. 


POLLINATION. Rinnex is wind-pollinated, the stigmas are therefore large and 
brush-like (indicated in Fig. 354 B). llliciim and Polygonumaxe insect-pollinated 
and Lave therefore capitate stigmas, etc.; honey-glands are situated at the base 
of the stamens (d, in Fig. 35-1 C, and n in Fig. 356) ; a few small-flowered 
Polygomim species are self-pollinated ; Buckwheat (P. fagopyrnm) is dimorphic 
and has long-styled and short-styled flowers (Fig. S56). Pol. bistorta is protan- 
drous and homostyled. 

About 750 species, most of which are found in the temperate regions of the 
Northern Hemisphere, some reaching as far as the snow line or into the Arctic 
regions (O.ryriu, Klmigiu). Trees and shrubs are found in the Tropics: 
Coccoloba, Triplaris. Rheum is Central Asiatic. The thick rhizomes of li. 
officinali' (Rhubarb) are officinal. The rhizomes of the ordinarily cultivated 
species, I\. undulatum and r Impolitic urn, are used in veterinary medicine. The 
following are cultivated as culinary plants for the sake of their leaves : Rninex 
acetosn (Sorrel), 11. patient ia, R. scntattts, and Hheiiin itnditlatuut (petioles). 
Several species of Pulyyoniun (P. Injdropiper and others) have a sharp, pungent 
tastp. "Buckwheat " is the mealy fruit of Pulygonum fagopyrum (Central Asia) 
and is of value as a farinaceous food. P. cn*pidatum (P. sieboldi, Japan) is an 
ornamental plant. Calligouum in sandy and stony deserts. 

FIG. 356. Flower of Polygonum fagopyrnm in longitudinal section: 1, long-styled ; 2, 
short-ityleil ; a the anthers ; s( the stigmas ; n nectary. 

Order 2. Piperaceae (Peppers). Shrubs or herbs, often 
with nodose, jointed stem ; leaves simple, entire, often with curved 
veins ; stipules wanting (Peperomid) or intrapetiolar and cap- 
like, often enclosing- the terminal buds (Piper}. The flowers in 
the group Pipere<.e (Piper, Fig. 357, and Peperomin) are borne in 
spikes with fleshy axes (club-like), seldom in racemes, the outer 
ones are crowded and are ^ or unisexual, always small, naked and 
without bracteoles; generally stamens 3+3, and gynceceum 3, but the 
number of the stamens may be reduced by suppression to 2, and the carpels to 1. 
The flowers are situated in the axils of the small, generally 
shield-like floral-leaves. The ovary is always unilocular and has 
one upright, orthotropons ovule. Fruit a berry or drupe. Both 
endosperm and perixperm are present, the latter being especially 
well developed (Fig. 359). 

Piper ; generally shrubs with scattered leaves, and terminal 



inflorescences which, are crowded to one side by the development 
of the highest lateral bud, so that they are situated opposite tho* 
leaves (Fig. 357). Many species have stems with an abnormal 
anatomical structure. Peperotnia ; chiefly succulent herbs, often 
epiphytes, with opposite or verticillate leaves having aqueous 
tissue on the upper side. 

The group Saururea (considered by some as an order, and perhaps representing 
a more original type) has 3-4 carpels with many ovules. Lactoris stands the 
highest with regular 3-merous perianth, 3 + 3 stamens and 3 carpels, which are 

FIG. 357.- Piper nigrum : branch with fruit (\). 

united at the base. Fruit a capsule with several seeds. (It has one species 
from the island of Juan Fernandez, and is also placed in an order of its own, 
Lactoridaceas, allied to the Magnoliaceas, through Drimys). Sanrurus has naked 
flowers ; most frequently 6 stamens, and 4 carpels, free or united at the base, 
each with 2-4 orthotropous ovules. Fruit, small berries. Houttuyniu ; stamens 
situated a little upward on the ovaries ; placentation parietal ; capsule many- 

About 1,000 species ; entirely tropical, especially from South America and East 
India. They are found chiefly among the underwood in damp, shady places ; 
some, which are fleshy (Peperontia), live as epiphytes on trees; a few climb by 



roots. USES. Several Piperaceae are used medicinally arid for spices on ac- 
count of their pungent properties and the essential oils found in nearly all 
parts of the plant. The following are officinal: "Black-pepper" (the unripe, 
dried fruits) and "White-pepper" (the seeds of the ripe fruits) of Piper nigrum 
(climbing shrub, East Indian) ; " Cubeb " berries of P. cubeba (climldng shrub, 
Java. "Long-pepper" is the unripe inflorescence of P. longum, East India. 

FIG. 358. Piper nigrutn (Diagram). In 
addition to the bract there are two struc- 
tures resembling bracteoles. 


FIG. 359. Piper nigruni : Fruit in longi- 
tudinal section, showing the endosperm, 
perisperm, and pericarp. 

The leaves of P. angmtifolia (Matico) are officinal. The leaves of the Betel- 
pepper (East India) are used together with the nuts of the Areca-palm to form 
the well-known East Indian intoxicating compound "Betel." A good many 
others are also used. 

Order 3. Chloranthaceae. (Chloranthus, Hedyosmum) have opposite leaves, 
with stipules more or less united at the base, and inferior " drupes." Ovules 
pendulous. Only endosperm. About 33 species, Tropical. 

Family 7. Curvembryae. 

The plants in this family have a curved ovule, and most fre- 
quently a kidney-shaped seed (generally provided with tine, cuticular, 
projecting- warts, Fig. 362J>), with a curved, peripheral embryo en- 
closing the endosperm which is most frequently floury (Figs. 362 C, 
365 H; for exceptions, see Fig. 366) ; the seeds in all cases are 
borne on a centrally -placed, and in most cases free, placenta (they 
are " basal " when there is only 1 ovule in the ovary, Fig. 364). 
The flower is regular, hypogynous or perigynous (Fig. 364) (only 
rarely epigynous) and usually 5-merous. The flower which is most 
complete has 5 whorls (S5, P5, A5 + 5, G2 3 5), as in some 
genera of the Caryophyllaceae (Figs. 360, 361) ; but from this 
type it becomes reduced, the petals and stamens being suppressed, 
so that finally 5 perianth-leaves, 5 stamens (opposite the perianth- 
leaves), and 2 carpels (Fig. 361 F) only are present ; for example, 


in certain genera of the Canjophyllacete, in the Chenopodiacete, 
Amarantaceae, and others. When the number of stamens is in- 
creased to more than 5 in the whorl, it is always possible to show 
that some of the stamens have been divided. The number of the 
carpels and ovules also becomes reduced ; in the highest there is a 
central placenta, riot free in its early stages, with a large number 
of ovules ; in those which are most reduced there is only a single 
ovule, which is placed centrally at the base of the ovary (Fig. 364). 
Somewhat corresponding changes are found in the fruit, which is a 
many-seeded capsule in those which have many ovules, but a one- 
seeded nut where there is one ovule. In the most reduced forms 
the flowers are generally unisexual. Similar features are also pre- 
sent in the vegetative parts. Almost all the species are herba- 
ceous, the leaves are simple and most frequently without stipules. 
The structure of the stein, especially in Chenopodiaces, Amarantacese, Nyctagi- 
niaceae and others, often differs from that of the ordinary Dicotyledon. In the 
woody portion of the stem and root several rings are sometimes formed which 
resemble annual rings but which are formed by new cambium-rings arising out- 
tide the old ones which then cease to divide. 

Order 1. Caryophyllaceae. Herbaceous plants, with round, 
nodose stem ; leaves opposite, slightly amplexicaul, simple, with 
sessile, undivided, entire lamina; stipules nearly always absent ; the 
inflorescences are dichasia passing over into unipared scorpioid 
cymes. The flowers are regular, $ or unisexual, hypogynous ov 
perigynous,5- (or 4-) merous with 2-3-4-5 carpels; calyx persistent ; 
corolla polypetalous. The ovaiy is unilocular (or originally, and 
sometimes also in the later stages, plurilocular below, e.g. Viscaria), 
with free styles and 1-several curved ovules on a central, free 
placenta. The fruit is a nut or a capsule opening apically with 
long or short valves (teeth, Fig. 362J, equal to or double the carpels. 
For the seeds refer to the family. In Dianthus the embryo is straight. 

The flowers which are most complete have Sn, PH, An + u (obdiplostemo- 
nous), Gn where n = 5 (Figs. 300, 3bl A) or 4 (Fig. 361 B) ; the carpels may be 
placed opposite to the sepals (Fig. 360) or opposite to the petals (Fig. 361 A, B). 
Without any change taking place in the position of the other whorls, the 
carpels are next found reduced to 2-3-4 (see the geutra) ; their number may 
easily be recognised by that of the styles. This is the construction in the 
majority of the genera in the two first groups. Stelluria media differs consider- 
ably. It may have (u) the flower as described above, with G6 ; (b), the corolla 
only absent, or (c) only the petal-stamens (Ao + 0, Fig. 361 C), or (d) all these as 
well as some of the sepal-stamens. The same applies to Saying, Alsine, Ccras- 
tiuin, and others, and, finally, a series of genera are formed, with certain coudi- 



tions of reduction which have become constant, and by a gralual series of steps 
lead to the most reduced form, which has only 5 sepals and 5 (or even as far as 
only 1) sepal-stamens (Fig. 361 D, E, F). The 
PETALS in the Ahinea; are often deeply bind. The 
sepal-stamens are most frequently the longest, and 
bear nectaries at the base (Fig. 363 st}. In the 
most complete forms the ovary has partition-walls 
in the lower portion (Fig. 360) ; these do not, 
however, reach to the top, and generally soon dis- 
appear. The ovules, when numerous, are situated 
on the placenta in as many double rows as there 
are carpels. In the number of ovules a reduction 
from many to 1 takes place (Fig. 361). A com- 
parison proves that the " free, centrally placed " 
placenta is formed by the vental portion of the carpels. The single basal ovule 
in Herniaria (Fig. 364), Scleranthns, and others, is also borne on the carpels. 

The vegetative branching is characteristic. One of the leaves in a pair is 
formed before the other, and has a more vigorous axillary bud ; these stronger 
leaves stand iu a ^-spiral, the fifth above the first one, and the branches are 
consequently arranged in the same manner. In the inflorescence, however, it 
is the upper or second bracteole 1/3) whose axillary bud (10 in Fig. 361) is most 
advanced. The bud of the first bracteole (a) bacomes sometimes entirely sup- 
pressed, or in some this bracteole itself is supprassed. 

FIG. 360. Diagram of Lych- 
nis : a, ft bracteoles. 

PIG. 361. A-F Diagrams of flowers of the Caryophyllacere : A Agrostemma ; B Sagina ; 
C Stellaria- D Corrigiola; E Paronycliia; F Herniaria. 

The most original type appears to be represented by the Alsiuefe. From this 
form on one side the Sileneae, adapted in a higher degree for insect-pollina- 
tion, are developed, and on the other side the Paronychieae, with various re- 

I. ALSINE&, STITCHWORT GROUP. Sepals free, and connected 
with them stellately expanded, slightly iinguiculate (white or in- 



odorous) petals; these, however, often become suppressed (Fig. 
363). The fruit is a capsule. 

a. As many carpels as sepals (4 or 5). Cerastium (Chick- 
weed). The petals are bifid. Capsule cylindrical, frequently 
curved at the top, and opening by 10 teeth (Fig. 362). Malachium 

A B C 

FIG. 362. Cerastium arvense : A fruit; B seed; C section of seed. 

differs only in the 5-toothed capsule with bifid teeth. Spergula (Spurry). 
The petals are not bifid , capsule 5-valved ; seeds winged. The 
leaves are linear, and appear as if placed in large numbers in a 
whorl, a branch being situated in the axil of each with leaves 
placed very close together at its base ; stipules membranous. Sagina 
has Sn, Pn, An + n, or An, Gn, where n=4 or 5. The corolla is often wanting. 

b. 3 (rarely 2) carpels (Fig. 361 C). Stellaria (Stitchwort) 
has deeply cleft petals. The number of stamens varies (see above). 

FIG. 363. Arenaria (Halianllius) peploides : (A) and <J -flower (B, C). 

Arenaria has entire petals. (To this group belong Alaine, Moehringia, 
Halianthus, or Honckenya (Fig. 363), which differ from each other, especially in 
the form of the seed and number of the capsular valves.) Spergularia has 
membranous stipules, as in Spergula. Holosteujn. 

2. PAEONYCHIE^E (Figs. 361 D, E, F; 364). Small, greenish 


plants. The loaves, in the majority, are opposite, with membran- 
ous stipules. The flowers are most frequently arranged in small 
dicha.ria ; they are small and insignificant, perigynous (Fig. 364) 
or hypogynous. The corolla is in most cases wanting, and when 
present is A^ery small ; in general the calyx-stamens are developed, 
but the corolla-stamens may be represented by small scales (Fig. 
364). Ovary most frequently with 1 ovule. Fruit, a nut, rarely a 
capsule ; it is enclosed by the strongly perigynous floral axis 

Scleranthus (Knapwell) is perigynous with bell-shaped torus ; 
no corolla; corolla-stamens are wanting or rudimentary; some 
calyx-stamens may also be absent. Corrigiola (Fig. 361 D) ; 
Illecebrum ; Paronychi'a (Fig. 361 E) ; Herniaria (Figs. 361 F, 364). 

This has a gamosepalous calyx and un- 
guiculate, white or red, petals, with out- 
growths (ligide, corona, paracorolla} at the 
throat of the corolla. These structures 
are not found in the other groups, and 
are merely outgrowths at the junction of 
the limb and claw. The corolla, stamens 
and ovary are frequently raised above the 
calyx, upon a lengthened internode (gyno- 
pliore). The flower has S5. Po, A5 + 5 ; 
fruit a capsule with many seeds. 

a. 5- (rarely 3-4) carpellate ovary. 
Lychnis (Campion, Fig. 360). The corolla 
is longer than the calyx ; corona present. 
The capsule is 10- or 5-toothed, completely 
1- chambered or 5 -chambered at the 
base, the genus has been divided accord- 
ingly into several genera : Melandrium 

Lychnis, Viscaria. Some species are unisexual by the abortiou of stamens 
or carpels (L. vetpertina, diurna). Agrostemma (A. githago, Corn-cockle, 
Fig. 351 A) has a long-toothed calyx, the teeth exceeding the 
corolla ; corona absent; 5-toothed capsule. 

b. Tricarpellate. Silene (Catch-fly). Six-toothed capsule ; 

corona present in the majority. Cucubalus has berry-like fruits which 
finally become dry but do not dehisce. 

c. Bicarpellate (2 styles, 4-toothed capsule). Dianthus (Pink) ; 
at the base of the calyx 1-several pairs of floral -leaves are situated ; 

Fts. 361. Herniaria glabra : 
o 1 flower ; b 1 longitudinal sec- 
tion through the flower; c 1 stig- 
ma with two pollen-grains. 


corona absent. The straight embryo is a pecu'iar exception. Gypsophila 
lias a campanulate, open calyx, 5-nerved, membranous between the 
nerves; corona absent; the flowers are generally small and 
numerous, in a large, paniculate dichasia. Saponaria (Soapwort) 
has corona. 

POLLINATION. Alsinea has ordinary nectaries at the base of the calyx- 
stamens (Fig. 336) : they are frequently protandrous but may often, in the 
absence of cross-pollination (in the less conspicuous specie*) pollinate them- 
selves. Their open flowers are accessible to many kinds of insects (particu- 
larly flies and bees). Gynodioecious flowers are found in several species, and 
the 5 -flowers are then generally more conspicuous than the ? -flowers. That 
the "? -flowers have descended from $ -flowers is seen by the large staminodes 
found in them (Fig. 363). Arenaria peploides is dioecious (Fig. 363). The Sileiifie 
are as a rule adapted for pollination by insects with long probosces especially 
butterflies, and they are frequently protandrous, so that at first the cahx- 
stamens open, later on the corolla-stamens, then the stigmas expand. The 
honey is secreted by a ring-like nectary round the base of the ovary or by 
nectaries at the base of the stamens. Some only blossom and emit scent at 
ni^'ht or in the evening (Lyd.nis vespertina, Silene initans, Sajionaria otfici)ialin) 
and, like other night-flowers, are of a white or pale colour. 

DISTRIBUTION. 1,100 species, especially in temperate climates, fewer in the 
colder zone, less still in the Tropics. The Paronychieae are especially found in 
dry, sandy fields. 

USES. " Soap-root " (with Saponiii, forming a lather in water) from Saponaria 
officinalis was formerly officinal, and GypsopJiila stnitlihnn. The seeds of 
Aaro*tenima githago are said to be poisonous. The following are ornamental 
plants : species of Pinks (D. caryophyllus, garden Pink, often with double 
flowers; D. barbatus, plumarius, etc). Lychnis, Gypfophila, Silene, Ceraxtitnn 
(C. tomcntosum as edging for borders), Saponaria officinalis (often coronate). 
Xpergitla arrenais is sometimes cultivated. 

Order 2. Amarantaceae. The flowers are essentially the same as in the 
Chenopodiacefc and the extremely reduced Caryophyllaceae (Fig. 361 F) ; they 
are regular, hypogynous, generally $ , have 5 free (rarely slightly united) 
] erianth-leaves ; in front of these 5 stamens, which are often united at their 
base into a shorter or longer tube and have stipule-like teeth between them (the 
division Gumphrfneteba-s 2-locular anthers, each of which opens longitudinally) ; 
and a 2-3 carpellate gynoeceum with one loculus and most frequently one, more 
rarely several ovules; the fruit is a nut, more rarely (in Celoxia, Ainarantnx, 
Gomphrena) a capsule, dehiscing irregularly, or like a pyxidium. The characters 
which especially separate them from the allied orders are found in the perianth. 
The perianth-leaves are not green and herbaceous, but inembranoun, dry, and 
often coli ured ; they are frequently produced into a bristle or awn ; they have 
also both subtending floral-leaves and 2 large bracteoles similar to the perianth ; 
all these dry leaves persist without alteration after the withering of the flower. 
The flowers are without scent. They are arranged in spike- or capitulum- 
like inflorescences ; sometimes placed singly, sometimes aggregated iu the 
panicle-like inflorescences ; in others, on the contrary, in dichasia. The 


majority are herbs, some are shrubs. The leaves are scattered, or opposite, 
but always simple and without stipules ; some are smooth, others hairy. 

450 species ; especially in the Tropics, principally S. Am. and E. Ind. : 
few are found outside these countries. Only a few are used; some, chiefly 
E. Indian species, are cultivated as ornamental plants: Amaranthus (Fox- 
tail) ; Gomphrena globosa ; Celosia cristata (Cock's-comb) remarkable for 
its fasciated inflorescence ; Alter nanthera. Some are employed as culinary 
plants in the Tropics, and in a few of the E. Indian species the seeds are 
farinaceous, and used for food. 

Order 3. Chenopodiacese. Generally herbaceous plants like 
the Caryophyllacefp, but the leaves are arranged spirally (except 
Salicornia), and are simple, exstipulate ; they are generally fleshy 
and like the stem " mealy," that is, covered with small hairs, whose 
large spherical terminal cell readily falls away; otherwise they 
are seldom hairy. The inflorescences are generally flower-clusters 
borne in panicles. Bracteoles generally absent. Flowers gene- 
rally unisexual : with the single exception of Seta the flowers are 
hypogynous; they are regular, small and inconspicuous, with single, 
green, 5-leaved, but more or less united perianth ; 5 stamens opposite 
the perianth, and a "2-5-carpellate, wiilocular ovary with 1 basal, 
curved ovule; but in some genera the number of the perianth-leaves 
and stamens is reduced to 3-2-1-0. The fruit is generally a 
nut, thus this flower and fruit are the same as in the reduced 
Caryophyllaeere (Fig. 361 F). The seed is similar to that generally 
found in the family (for exceptions see the genera). 

The floral diagram most frequently present is the same as in Fig. 361 F. There 
is no indication of corolla or of corolla-stamens, which may be supposed to have 
belonged to the plant, but which are now entirely and completely suppressed. 
This order appears to have been an offshoot from the Caryophyllacete. The 
perianth persists after the withering of the flower, and envelopes the nut ; it is 
very variable, and, together with the position of the seed, the form of the 
embryo, the sex of the flowers, etc., gives the characters of the genera. 

I. CHENOPODIEJ;, GOOSEFOOT GROUP (Fig. 365), has (or poly- 
gamous) flowers, with regular 5-parted perianth (G) ; the em- 
bryo is ring-like (-ZT). The leaves have the ordinary flat forms. 
Chenopodium (Goosefoot). The flower is hypogynous, and the 
fruit (which is compressed) perfectly free ; Mulberry-like collec- 
tions of fruits are formed in some species (sub-genus Blitum) by 
the perianth becoming finally fleshy and coloured. Beta (Beet, 
Mangold, Fig. 365) differs from all genera in the perianth, which 
finally becomes cartilaginous, being epigynous (D). Small, most 
frequently 2-3-flowered clusters without bracteoles, situated in a 

w. B. B B 



long, interrupted axis (A, -Z?) ; the flowers and fruits in each 
cluster are more or less united individually, and fall off together 

FIG. 365. Beta vulgaris. 

they are commonly known as seeds (E, F). The seed lies horizon- 
tally. Hablitzia (H. tamnoides). 

2. SALSOLE^, SALTWORT GROUP, has cylindrical or semi-cylindi-i- 
cal leaves. Perianth as in the preceding group ; the fruit is most 

FIG. 306. -Salsola soda: embryo. 

frequently compressed. The two first mentioned genera differ 
from most of the others in the order in having a spirally-coiled, 


and not a ring-like embryo, so that the endosperm is slight or 
wanting (Fig. 366). These plants are sometimes placed as a 
group by themselves, SPIROLOBE^ in contradistinction to which 
the others are termed CYCLOLOBEJ:. Salsola (Saltwort) ; leaves 
subulate, with spiny tips ; the flowers have 2 spinous bracteoles : 
during the ripening of the fruit a tough leathery wing is de- 
veloped transversely to the back of the perianth. Chenopodina 
deviates from Chenopodium chiefly in the embryo and want of endosperm. 
Koclria has a somewhat similar perianth to Salsola, but a ring-like embryo; 
it differs from the others in being hairy. 

3. SALICORNIE.E, GLASSWORT GROUP. Salicornia (Glasswort) has 
a very different appearance. The stems are succulent, jointed, and 
almost leafless ; the leaves opposite, very small, sheath-like and 
connate ; there is a depression in the axil of each leaf, in which a 
small 3-flowered dtchasium without bracteoles is sunk ; the 
flowers have a trimerous perianth, 1-2 stamens and 1 carpel. No 
endosperm. S. herbacea on clayey beaches. 

4. ATRIPLICE.E. This group has most frequently unisexual 
flowers ; the J 1 -flower has a 4-5 partite perianth, but the ? -flower 
differs from it. Atriplex is monoecious or polygamous, the $ - 
flower is naked, but has 2 large, herbaceous bracteoles which 
expand during the ripening of the fruit, and often become warted 
and fringed, enveloping the compressed nut. The section Dichospennnm 
has two kinds of $ -flowers, one like those just described, the other similar to the 
Clienoi>odiiim-&o\vers, which have been deprived of their stamens, and the fruits 
of which are depressed, not pressed together from the sides ; some (e.g. A. 
hortensis) have even three kinds of nuts. All the flowers of Atriplex, which 
present vertical fruits, are accessory shoots, which stand beneath the ordinary 
flower-clusters, a rather singular relation. Spinacia (Spinach) is dioecious ; 
( -flower: perianth, 4 (-5) ; stamens, 4 (-5) ; ? -flower: tubular, 
2-4-partite perianth, hardening during the ripening of the fruit, 
and uniting with the compressed nut ; in S. oleracea, it also forms 
thorns; 4 long stigmas. Halimus has the 2 long bracteoles 
almost entirely united and ultimately adhering firmly to the 

5. BASELLE.B. A somewhat exceptional group with more or less perigynous 
flowers and 2 bracteoles. Hasella, Bomsingaultia, Ullucus. The perianth is 
sepaloid ; ovary 1-ovuled. In Basella the perianth is fleshy, enveloping the 
nut, and the cotyledons are so rolled together that a tranverse cut divides them 
in two places (as in Spirolobeas). Herbaceous climbing plants. 

POLLINATION. "Wind- and self-pollination, as far as is known ; the insignifi- 
cant flovveis, devoid of honey, appear to exclude insect-pollination. 520 
species. Most of them are annual (out of 26 native species only 5 are peren- 


iiial) ; inhabiting salt-marshes and salt-steppes, and growing as weeds (most 
frequently on garden or field soil containing manure) in this country, especially 
species of Chenopodium and Atriplex. The majority are found outside the 
Tropics, and play a very important part, for example, in the Asiatic salt- 
steppes. They grow gregariously in large masses. 

USES. Comparatively few. The only important one is Beta vulgaris (from 
the Mediterranean basin), with its different varieties, viz. Beet-root, Cattle-beet 
or Eed-beet, Sugar-beet, and others. These are biennial, making in the first 
year a root which acts as a reservoir of reserve material, with a rosette of leaves, 
and in the second year using this material in the production of a long stem, 
leaves and flowers. The primary root has been developed by cultivation into a 
very thick and fleshy tap-root ; its mode of increase in thickness deviates from 
that of other roots, concentric rings of vascular bundles being formed from a 
carnbial ring developed outside the previous ring. In this way several rings 
of vascular bundles separated by medullary rays, alternating with rings of 
parenchyma, may be found in the root of a Beet. Besides Beta vulgaris, var. 
horten*is (Beet-root), the following are also cultivated : var. ciclu (Leaf-beet, 
"Mangold," or " Eoman Spinach "), Spinacia oleracea and Atiiplex liortemis 
as Spinach ; a form of the latter and of Spinach are grown as ornamental 
plants. The tubers of Ullucits tuberosus are used as potatoes ; Chenopodium 
quinoa, in Chili and Peru, is an important farinaceous plant. Soda is made 
from some (Salsola kali, Chenopodina maritima and others). Aromatic proper- 
ties are rare : Chenopodium ambrosioides and botrys. 

Order 4. Batidaceae. Batis maritima, a bushy West Indian maritime 

Order 5. Phytolaccaceae. The (sometimes unisexual), regular, some- 
times slightly perigynous flowers are inconspicuous and have a single sepaloid 
or coloured 4-5-leaved perianth (generally united at the base) ; stamens either 
in 1 whorl in the spaces between the perianth-leaves or in 1 whorl opposite the 
perianth-leaves, or in 2, one of which alternates with these ; but the number 
may be increased by the splitting of one or of both the whorls to as many as 
10-15-20-25. Carpels sometimes only one, sometimes many (4-10) placed in a 
whorl, either free or united into a gynceceum with a corresponding number of 
loculi in the ovary ; but in all cases each carpel bears only its own style and 
1 ovule. The fruit is a berry (or nut, capsule, or schizocarp). Mostly herbs 
or herbaceous shrubs, with scattered, simple leaves without stipules (PetiveriecB 
have stipules). Inflorescences, most frequently racemes or spikes, which in 
some instances are apparently placed opposite to a leaf, being displaced by a 
more vigorous growth of the axillary bud. Embryo always bent. Petiveria 
has a straight embryo with rolled cotyledons. Pliytolaeca, Pircunia, Microtea, 
Seguieiia, Rivina (Pr4, A4. Gl; berry), and others. 

The following plant is, with some doubt, placed near this order : Thely- 
(jonum cynocrambe ; monoecious. $ -flowers: perianth, 2-leaved; stamens indefi- 
nite. <j? -flowers : perianth-leaves united, 3-toothed ; Gl, style gynobasic. 
Fruit a drupe. An annual plant ; Mediterranean. Branching anomalous. 

About 90 species ; in tropical and temperate countries, principally America 
and Africa. The red juice in the fruits, especially of Phytol. decandra, is used 
for colour'ng wine. 


Ch-der G. Portulacacese (Portulacas). The flowers are 
regular (except Montia), hypogynous (except Portulaca) and $ 
The diagram which applies to the majority of genei-a is that in 
Fig. 307, but with all the 5 stamens completely developed : it may 
be considered as the Chenopodiaceous diagram with the addition 
of 2 bracteoles in the median line (m-n, these by some are con- 
sidered as sepals), and with a petaloid perianth (usually desig- 
nated "corolla"). The "petals" fall off very quickly, and are 
sometimes wanting. Most frequently 5 stamens, situated oppo- 
site the " petals," but in other genera the number varies ; Mcntta 
has only 3 stamens (by suppression of the two anterior and 
lateral, Fig. 367), others again have more than 5, some a large 
and indefinite number. This may be explained partly by the 
appearance of a second whorl of stamens alternating with the 
first, and partly by the splitting (dedoublemeiit) of the stamens. 
Grynceceum most frequently tricarpellate, ovary unilocular with 
1-several basal ovules (sometimes on a 
branched placenta, as in certain Garyo- 
phyllacece). The fruit is a capsule, more 
rarely a nut. The majority are annual 
herbaceous plants with scattered, entire 
leaves, often fleshy and smooth, with or 

without rudimentary stipules (dry, mem- 

J FIG. 367. Montia. 

branous, modified into hairs). Inflorescence Diagram of flower, 


Portulaca (Portulaca) : flower, epigynous or semi-epigynous ; 
fruit, a pyxidium. The stamens vary in number, and are most 
frequently placed in groups (in consequence of splitting) opposite 
the petals. Montia : the corolla is slightly gamopetalous, but 
cleft on the posterior side (Fig. 367), and as a consequence of the 
larger size of the lateral petals, slightly zygomorphic ; 3 stamens. 
Calandrinia ; Talinum ; Anacampseros ; Claytonia. 

125 species ; mostly in warm and temperate countries, especially the arid parts 
of S. Am. and the Cape. Montia fontana (Blinks) is a native plant. Portulaca 
oleracea is cultivated as a pot-herb in the south of Europe. A few species of 
Portulaca and Calandrinia are ornamental plants. 

Order 7. Nyctaginiaceae. The characteristic feature of this 
order is the single, regular, united, and often petaloid perianth, the 
lower part of which generally persists after flowering and em- 
braces the fruit as a false pericarp. The upper portion is most 


frequently ralrate and folded, or simply valvate in Aestivation. 
The number of stamens varies. The free gynreceum is unicar- 
peUate and has 1 ovule. The fruit is a nut, but becomes a false 
drupe, since the lower persistent portion of the perianth becomes 
fleshy (as in Neea, where this fleshy part is almost always crowned 
by the upper persistent part of the perianth. In the majority of 
the Mirabilese the lower part becomes the dry anthocarp, while 
the upper petaloicl part falls away after flowering). Finally, a 
peculiar involucre is formed around the flowers by free or 
united floral-leaves. -The majority are herbs, some are trees 
(Pisonia, etc.) ; Bougainvillea is a liane. The stems are often 
nodose and swollen at the nodes; the leaves are simple, penni- 
nerved, scattered, or opposite, without stipules. In some, the vascular 
bundles are scattered ; stem anomalous. 

Mirdbilis ; the structure of the stem is abnormal. Dichasial 
branching with continuation from the second bracteole, thus form- 
ing uni pared scorpioid cymes. The perianth is petaloid, funnel- 
shaped, and has a folded and twisted aestivation resembling that 
of the corolla of the ConvoLrulacece; the upper coloured portion 
falls off after the flowering. Outside, and alternating with it, is 
a 5-partite, sepaloid involucre of 5 spirally-placed floral-leaves. 
OxybapJius; the involucre envelops 1-3 dichasial flowers. - 
Bougainvillea; the involucre is rose-coloured, 3-leaved, and en- 
velops 3 flowers (placed laterally; the terminal flower wanting). 
The leaves of the involucre in Boerhaavia, Pisonia, Neea, and others 
are reduced to teeth or scales. 

157 species; mostly in tropical countries, and especially S. Am. Species 
of Mirabilis (Am.) are ornamental plants. Thein is found in Neea thelfera 
Oersted (discovered by Lund in Lagoa Santa, Brazil), which may be used as a 

Order 8. Aizoaceae. Only 3 whorls are found in the flower, which alternate 
with one another when their leaves are equal in number. The first is sepaloid, 
the th ; rd one the carpels, and the intervening one is either uncleft, in which 
case it is developed as stamens, or it is divided into a large number of members 
which then all become stamens (arranged in groups), or the outermost ones 
become developed as petals. The fruit is most frequently a capsule with 
several locnli. Most of the species are herbs with thick, fleshy stems, and 
exstipulate leaves. The structure of the stem is usually anomalous. 

i. AIZOIDEJE have hypogynous or perigynous flowers with (4-) 5 perianth- 
leaves ; stamens single, or (by splitting) in groups of 2-3, alternating with the 
perianth-leaves. The gynoaceum (with 3-5 carpels) has 3-5 loculi in the ovary, 
and most frequently numerous ovules in each loculus, borne on the central 
placenta formed by the edges of the carpels. The fruit is a capsule. The 


inflorescences are diohasia and unipared scorpioid cymes. Aizoon, Mollujo, 
Sesuviian, and others are herbs or bushes, most frequently hairy. 

2. MESEMBRIANTHEME.E have senii- or wholly-epigynous flowers. Tetra- 
gonia. The perianth is 4 (more rarely 3-5-6) -merous. Stamens single, or (by 
splitting) in groups alternating with the perianth- leaves. There is an indefinite 
number of carpels, and each loculus of the ovary contains only 1 pendulous 
ovule. Fruit a nut or drupe. The flowers arise singly in the leaf-axils, with an 
accessory foliage-bud below them ; in some instances there is also an accessory 
flower between this bud and the flower. Southern hemisphere, especially at 
the Cape; T. e.rpunsa, New Zealand Spinach, is a fleshy plant which is 
cultivated as a pot-herb (Japan, Austr., S. A.m.).Mesembrianthemum : the 
flowers are 5-merous ; the numerous linear petals and the still more nu- 
merous stamens all arise by the splitting of 5 or 4 protuberances (primordia) 
alternating with the sepals. The ovary presents another characteristic peculi- 
arity : the carpels alternating with the 5-4 stamens form an ovary (with several 
loculi) with the ovules at first borne, as in other cases, on the inner corner 
of the inwardly-turned carpels ; but during the subsequent development the 
whole ovary is so turned round that the placenta become parietal and the 
ovules assume, apparently, a position very rarely met with in the vegetable 
kingdom : on the dorsal suture of the carpels. Shrubs or under-shrubs, more 
rarely herbs with fleshy stems and simple, entire, more frequently thick or 
triangular leaves, containing a quantity of water. The flowers open about noon, 
and are brightly coloured, generally red or red-violet, but odourless. The 
capsules dehisce in rainy weather. 300 species, mostly found at the Cape. 
Some are ornamental plants. M. crystalliinuii (the Ice-plant) and others are 
covered with peculiar, bladder-like, sparkling hairs, the cell-sap of which con- 
tains salt these serve as reservoirs of water. 

Family 8. Cactiflorae. 

The position of this family is very doubtful ; but it seems in 
many respects to approach Mesembrianthemum. Some botanists 
place it near to the Ribesiacese ; others, again, to the Passinoracete. 
Only 1 order. 

Order Cactaceae (The Cacti). The flower is epigynous, $ , 
regular, and remarkable for its acyclic structure ; there are, for 
instance, a large number of spirally-placed sepals and petals, which 
gradually pass over into one another, and which in some species, to 
a certain extent, arise from the walls of the ovary as in Nymphssa 
(Fig. 383 A, I>). The petals are free; rotate, opening widely in 
Opunfia, Pereskia, and Ithipsalis ; erect and united at their base 
into a shorter or longer tube in Cereus, Epiphyllum, Mammillaria, 
Echinocactns, Melocactus, and others (Fig. 369). Stamens numerous, 
attached to the base of the corolla ; gynceceum formed of many 
carpels, with one style, dividing into a number of branches corres- 
ponding to the number of carpels ; the ovary has one loculus with 



many parietal placentae ; the ovules are anatropous, on long and 
curved funicles. Fruit a berry with exendospernious seeds. The 
fruit-pulp is mainly derived from the funicles. The external 


FIG. 363. A Echinocactus : o position of a leaf-lamina; b a lateral shoot on the displaced 
axillary bud. B Pereskia : b a foliage-leaf on a small thorny branch which is subtended 
by a foliage-leaf which has fallen off and left a scar(o). 

appearance of the Cactacea? is very peculiar; Pereskia, which has 
thick and fleshy leaves (Fig. 368), deviates the least ; foliage- 
leaves of the usual form are wanting in the other genera, or are 
usually very small, and quickly fall off and disappear (Opuntia), 
or are modified into thorns ; the stem, without normal foliage- 
leaves, so characteristic a feature in this order, makes its appear- 
ance after the two normally 
developed cotyledons. The 
stems are fleshy, perennial, 
and may finally become 
woody. In some they are 
elongated, globose, pointed, 
and more or less dichoto- 
mously branched, e.g. in 
several of the Rhipsalis 
species, which live mostly 
as epiphytes on trees ; in 
others, elongated, branched, 
globose, or, most frequently, 
more or less angular (pris- 
matic) or grooved and pro- 
vided with wings, and either 
columnar and erect (as 
much as about 20 metres in 
FIG. 369. Echinupsis. height and 1 metre in cir- 


cumference, as in C. giganteus in New Mexico) or climbing by roots 
(Cerens and Rhipsalis-species) ; in others again, compressed, more or 
less leaf-like, often with a ridge in the centre (winged), branched 
and jointed : Epiphyllum, Phyllocactus, Opuntia, some species of 
Rhipsalis ; others are thick, short, spherical or ovoid, unbranched or 
only slightly branched, and either studded with prominent warts 
(mammillce) each of which supports a tuft of thorns (Fig. 368 A 
Mammillaria and others) or with vertical ridges, separated by 
furrows (rows of mammilla^ which have coalesced) in Melocactus, 
Echinocactus, Echinopsis (Fig. 369) ; at the same time the ovary in 
some is embedded in the stem so that leaves or leaf-scars, with 
tufts of thorns in their axils, may be observed on the ovary just as 
on the stem. The flattened shoots of the Cactacese are formed 
in various ways, either by the compression of cylindrical axes 
(Opuntia) or, as in Melocactns, etc., from winged stems in which 
all the wings are suppressed except two. 

The tborns are produced directly from the growing points of the axillary 
buds, and are modified leaves. The axillary bud is united at its base with 
its subtending leaf, which as a rule is extremely rudimentary ; and these 
together form a kind of leaf-cushion, larger in some genera than iu others. 
This leaf-cushion attains its highest development in Mammilla ria, in which it 
is a large, conical wart (see Fig. 368 A), bearing on its apex the tuft of thorns 
and rudimentary lamina. The seedlings have normal cotyledons and a fleshy 

All the species (1,000?) are American (one epiphytic species of Rhipsalis is 
indigenous in S. Africa, Mauritius and Ceylon), especially from the tropical 
table-lands (Mexico, etc.). Some species, especially those without thorns, as 
lihipmlis, are epiphytes. Opuntia vnlrjaris, the fruits of which are edible, is 
naturalized in the Mediterranean. The cochineal insect (Coccus cacti} lives on 
this and some closely allied species (0. cocci nellifera, etc.), particularly in 
Mexico and the Canary Islands. Several are ornamental plants. 

Family 9. Polycarpicae. 

The flowers as a ride are 5 > regular and hypogynous ; however in 
some orders they are unisexual, e.g. in the Myristicacese, or 
zygomorphic (in Monkshood and Larkspur in the Ranunculacere) ; 
in the Lauraceee, (Fig. 386) for example, perigynous, and in 
NympJuea (Fig. 383) even partially epigynous flowers are typical. 
The flowers are acyclic in very many of the genera of the two first 
orders, if not completely so, at any rate in the numerous stamens 
and carpels, thus denoting an old type. It is a remarkable 
characteristic that in the majority of the orders the number 3 pre- 
vails in the calyx and corolla ; the number 5 also occurs, but the 



number 2 is seldom met with. Most orders have a double 
perianth ; chorisis does not occur, suppression is rare, and the parts 
of the flower are developed in acropetal succession. The most 
characteristic feature in the order is the free, one-leaved, as a rnle 
numerous carpels (apocarpous gynceceum). The number of carpels 
in some of the last mentioned orders dwindles down to 1 (e.g. the 
fjerberidex and Myristicacese) . The carpels in Nymphteacefv become 
united into one pistil (sjncarpous), a condition which we also find 
distributed among the other orders. 

Endosperm occurs in almost all the orders (except e.g. Lanracew). 
The nutritive tissue in Cabombefe and Nyiiiph&ete is chiefly peri- 

Order 1. Ranunculaceae Nearly all are herbs (except 
Clematis). The leaves are scattered (except Clematidefe), they 
have a large sheath with broad base (no stipules), and are most 

FIG. 370. Diagram 
of Aquilegia viilgaris : 
sj) spur. A cyclic 

FIG. 371. Diagram of a dichasium 
of Ranunculus ncer : a lt a 1 , and /3,, 
/3 1 , bracteoles (the buds in tlie axils 
of the bracteoles, a and a 1 , are con- 
tinued antidromously). The flower 
has cyclic calyx and corolla, but 
acyclic ( ? " T ) stamens. 

Fic.372. Diagram of an 

acyclic Ranunculaceous 
flower (only 3 stamens are 
indicated). The spiral of 
the sepals has a diver- 
gence of | ; that of the 
corolla and subsequent 

leaves f . 

frequently palminerved with palmate lobes. The flowers are 
hypogynous, with most frequently a well pronounced convex re- 
ceptacle (Figs. 374 7?, 380), $, regular (except Delphinium and 
Aconituni) ; their structure varies very much ; in some the leaves 
are verticillate, in others arranged spirally ; in others, again, both 
modes of arrangement are found. It is a characteristic feature 
that the various series of leaves (especially calyx and corolla) are 
not so distinct or so sharply divided as is usual. The leaves of the 
perianth are free, imbricate (except Clematidete} ; stamens numerous, 
with most frequently extrorse anthers; gynoaceum free, apocarpous 
(except NigeUa and partly Helleborus), with 1 or several ovules 
(Figs. 373, 378, 379) borne on the ventral suture. The fruit is 


either a nut or a follicle (Act&a has berries). The seed has a large, 
oil-containing endosperm and a small embryo (Fig. 374). 

The main axis generally terminates in a flower, and the lateral axes branch 
in a cyrnose manner (Fig. 371). The flowers show the following differences in 
constiuction: VEBTICILLATE (EUCYCLIC), i.e. constructed all through of alt mating 
whorls: Aqiiiletjia (Fig. 370), Xanthorhiza, and sometimes Erantliis. SEMI- 
YERTICILLATE (HEMicYCi.ic) /.c. with sepals and petals in alternate whorls, and 
the others arranged spirally: Ranunculus (Fig. 371), Mi/osiirus, Pteonia and 
several other genera entirely, or in certain species only. SPIRAL-FLOWERED 
(ACYCLIC) i.e. all the leaves are arranged spirally, so that sepals and petals do 
not alternate the one with the other, even though they are the same in number : 
Adonis (Fig. 372), Aconitinn, Delphinium-species, Nigella -species, Hellcborus. 
The leaves of the calyx are in this instance arranged on a spiral of | ; those of 
the corolla on f, f, ^ or 2 R T , and stamens and carpels likewise on higher 
fractions of the same series. 

The genera Caltlia, Anemone, TJialictrum and Clematis have a single perianth, 
which is most frequently petaloid ; it is thus apparent that the sepals are 
petaloid, and the leaves, which in other genera have developed as petals, are in 
these instances stamens. The calyx is similarly petaloid in the genera Hclleborus, 
Erantliis, Nigella, Delphinium and Aconitum ; but the petals are present in these 
instances in unusual (horn-like) forms, and almost entirely given up to the 
function of nectaries, a function they already possess in Ranunculus. According 
to a more recent theory the " honey -leaves " are transformed stamens, which 
have lost the function of reproduction ; the perianth is then single, and most 
frequently petaloid. [Those leaves in the flowers of many Ranunculac^ffi 
which bear nectaries are termed by Prantl honey-leaves, and comprise those 
leaf-structures of the flower whose essential function lies in the production 
<>f nectar, and which, independent of the differentiation of the perianth into 
calyx and corolla, are derived from the stamens by the loss of their reproductive 
functions. Clear transitional forms are found between the two series of the 
perianth (^.17. between the sepaloid and petaloid perianth-leaves of Anemone 
japonica, A. decapetalu, Troll 'ms-species) while transitional forms are never 
found between perianth- and honey-leaves (with the exception of Aquilenia 
vulgaru, var. stellata). In Anemone and Clematis the honey-leaves pass 
gradually into the stamens, and agree with the stamens in the other Rauuu- 
culaceae in their arrangement, development, and scant system of veins (except 
Nigella). In Delphinium, sect. Consolida, the two honey-leaves placed in front 
of the unpaired perianth-leaf are united into one, as shown by the veins (twice 
three veins arranged symmetrically). The honey-leaves of A quite a ia, Calliun- 
tliemnm, and the majority of the Ranunculus-species serve by reason of their 
large circumference, as organs of attraction, and on this account are considered 
as petals by other authors. The same position in the flower which the 
honey-leaves assume is found occupied by staminodes, without nectar, in some 
C >ptis-species, in Anemono2)sis, Acttca sect. Euactica, {e.g. A. racemosa), Clematis 
sect. Atrageue; in the last-named they closely surround the stamens, in Actcc.a, 
they are petaloid. A perianth, sharply differentiated into calyx and corolla, 
and destitute of honey-leaves, is found in Anemone, sect. Knoirltonia (Cape), 



Adonis, Pteonia. The perianth of the Ranunculaceffi is considered by Prantl 
to be usually petaloid. The nectaries arise in the Ranumulaceffi (1) on 
normal stamens (Clematis sect. Viorna), (2) on the h' ney-leaves (this is 
generally the case), and (3) on the carpels (Caltha and the majority of Trollius- 
species). As the result of his researches upon the Ranunculacese, Prantl 
does not agree with the view advanced by Drude (Schenk, Hand. d. Bot. iii.) 
that the petals in general have proceeded from the metamorphosis of the 
stamens (A')] . 


FIG. 373. Ovaries in longitudinal section : v the ventral suture; <i the dorsal suture : 
A, B Clematis ; C Ranunculus ; D Myosurus. 

The most primitive form of fruit is undoubtedly the pod formed by one carpel, 
on the edges of which (along the ventral suture) two rows of ovules are situated : 
Paeonieffi, Helleboreae, Delphiniea? (Fig. 379). In a great many genera the 
number of ovules has been limited to one perfect one, which is placed in the 
central plane under the united leaf-edges, and sometimes also some barren 
ovules above it (Fig. 373). The fruitlets in this case become achenes, and 
are present in much larger numbers than when there are follicles. 

FIG. 37-1. Hcllebonis niger : A flower; B receptacle ; pet petals PIG. 375. Caltha 

(honey-leaves); pi stamens and carpels; Cseed; D anther (cross palwsiris: frui*. 

section) ; <ilb endosperm. 

The following have FOLLICLES : Pxoniete, Hellebores (except 
Acttva) and Delpliiniex ; ACHENES : Ranuncideie, Anemonese and 



FIG. 376. Aquilegia vulgarls. 

A. Follicles (Figs. 375, 379), with many ovules, situated in 

two rows along the ventral suture. Actaa has berries, Niijella has 
capsules of several loculi. 

1. P^ONIE^E, PEONY GROUP. This has regular, acyclic flowers 
with a normal, most frequently 5-leavecl, imbricate calyx ; large, 
coloured petals, and 

introrse anthers. 

Slightly perigynous. 

Surrounding the 

base of the carpels 

a ring-like swelling 

of the receptacle 

("disc") is present, 

which is largest in 

P. m out an. The 

follicles are more 

or less fleshy or 

leathery. Mostly 

herbs, with pinnati- 

sect or decompound 

leaves and large, 

solitary flowers ; a gradual transition may be traced from the 

foliage-leaves to the petals. Pfeonia ; Hydraslis. 

2. HELLEBORES, HELLEBORE GROUP. This has regular flowers with 
most frequently a coloured 

calyx. The petals (honey- 
leaves) are modified into 
nectaries ; they may be horn- 
like, provided with a spur, or 
of a similarly unusual form, 
or they may be entirely ab- 
sent. Anthers often extrorse. 
Trollius (Giobe-flower 1 ). 
The flower is acyclic : many 

1 According to Prantl, some 
species of Trollius (T. enropams, 
and asiatiacus) have a perianth, 
differentiated into calyx and cor- 
olla, which does not pass over into 
the honey-leaves. The outer leaves 
of the perianth have frequently an 
incised apex, the intermediate ones FIG. 377 -CaK Tin palustns (nat. size). 


petaloid sepals, succeeding these, most frequently, several linear, 
dark yellow petals, which bear a naked nectary at the base; 
finally, many stamens and carpels arranged in a spiral (-, / T ). 
Caltha (Marsh-marigold, Figs. 375, 377); 5 (-7) yellow sepals, 
no petals. The foliage-leaves have a large amplexicaul sheath. 
Helleborous (Hellebore) has pedate leaves. The flower is acyclic, 
with 5 large, regular, persistent, often petaloid sepals (f) ; small, 
hornlike petals (honey-leaves ; most frequently 13, divergence -^3) 
and generally few carpels (Fig. 374). Coptis. Isopyrum. 
Eranthis (Winter Aconite), like Anemone, has a 3-leaved invo- 
lucre and most frequently trimerous flowers, 6 large petaloid sepals, 
6 petals (tubular honey-leaves), 6 oblique rows of stamens, 3-6 carpels. 
Aqnilef/ia (Columbine, Fig. 376) ; the flower is entirely cyclic 

FIG. 378. Nigella : A, B fruit of N. tlamnscena, entire, and cut transversely. C Petal 
(honey-leai) of N. arvensis. D Petal of N. damascena. 

and has large spurs on all the 5 petals (funnel-shaped honey- 
leaves); S5 coloured, P5, A5 x (8-12), G5 in regular alternation 
(Figs. 376, 370) ; the innermost stamens are often staminodes 
(Fig. 370). Nigella (Love-in-the-mist, Fig. 378) has 5 sepals 
and 8 small, two-lipped petals cleft at the apex (the nectary is 
covered by the under-lip; Fig. 378 C, D). The 5 carpels are 
more or less completely united; and a many-carpellate ovary 
with free styles is formed in some. Large air-chambers in the 
external wall of the ovary are formed in N. damascena (Fig. 378). 
Actfca (Baneberry) has coloured sepals, either no petals or an 

sometimes present transitional forms to the inner, and sometimes there is a 
distinct boundary between them. 



indefinite number, and only 1 carpel. The fruit is a berry (or 
follicle). Cimicifur/a, Gaiiddla, Xaitthorhiza (S5, P5, A5 + 5, G5). 

3. DELPHINIE^, LARKSPUR GROUP. Zygomorphic flowers witb 
coloured calyx ; the 2 posterior petals (honey-leaves) are trans- 
formed into nectaries, the others are small or absent altogether. 
Aconitum (Monkshood) ; 5 sepals, of which the posterior one (Fig. 
379 A) is helmet-shaped ; most frequently 8 petals (as in Fig. 372), 
of which the two posterior ones (honey-leaves) are developed into 
long-clawed nectaries (Fig. 379 .4, fc) enveloped by the helmet-like 
sepal; the others are small, or are to some extent suppressed. 
Stamens on a spiral of f-^ ; generally 3 carpels. Perennial herbs. 
Delphinium (Larkspur) ; very closely allied to Aconitum, but the 
anterior 4 petals are most frequently wanting, and the 2 posterior 
ones have each a spur, which 

is enclosed by the posterior 
sepal, the latter being also 
provided with a membranous 

spur. Stamens and carpels ar- 
ranged on a spiral of jf, -f^, ^. 
In P. njacis and consolida there is 
apparently only 1 petal (by the 
fusion of 4) and 1 carpel. 

B. Fruit achenes. Many 
carpels, each with only 1 
ascending (Fig. 373 C), or 
pendulous (Fig. 373 D), per- 
fect ovule ; often also rudi- 
mentary ovules above it (Fig. 
373 A, B). Fruit achenes. 

CUP GROUP, has double peri- 
anth. Hyosurus and Adonis have 
pendulous ovules as in Anemones 

(Fig. 373 >) ; Ranunculus, with Batrachium and Ficaria, erect ovules (Fig. 273 C) 
and downwardly-turned radicle. Ranunculus. Most frequently S5, P5, 
many spirally-placed stamens and carpels (Figs. 371, 380). The 
petals (honey-leaves) have a nectary at the base, covered by a small 
scale. Batrachium, Water Ranunculus, deviates by the acheues being trans- 
versely wrinkled ; dimorphic leaves. Ficaria has 3 sepals and 7-8 petals arranged 
in -. F. ranunculoides (the only species) has tuberous roots, which spring from 
the base of the axillary buds, and together with these, serve as organs of repro- 
duction. The embryo has only 1 cotyledon. Myosums (Mouse-tail) has 

FIG. 379. Aconitum napellus. A Flower in 
longitudinal section, below are the 2 bracteoles ; 
a half of helmet-like sepal ; b and c other 
sepals; fc nectary; / carpels. B Ovary in, 
longitudinal section ; C the same traLsversely ; 
d dorsal suture ; v ventral suture. 



small prolongations from the 5 sepals ; 5 narrow petals which bear 
the nectaries near the apex ; sometimes only 5 stamens, and an 

ultimately very long recep- 
tacle, with numerous spirally- 
arranged achenes (Fig. 381). 
Adonis is acyclic (Fig. 372) ; 
most frequently 5 sepals with 
a divergence of -f, 8 petals of 
, indefinite stamens and car- 
pels of | or f\. The corolla 
has no nectary. 

GROUP, has a single pei-ianth. 

(Pendulous ovules (Fig. 373 D), radicle turned upward) Anemone has a 

single, petaloid, most frequently 5-6-leaved perianth, and beneath 
the flower most frequently an involucre of 3 leaves, placed close 
together in the form of a whorl. In A. nemorosa, ranunculoides, 
etc., the involucral leaves resemble foliage-leaves ; in A. liepatica 

FIG. 380. Flower of Ranunculus sceleratus 
in longitudinal section. 


FIG. 381. Myosurus minimus: c cotyledons; m the foliage-leaves; f the floral axis 
with the carpels, and g the same without; y insertion of perianth. 

they are situated close under the perianth, and resemble sepals, 
and in the sub-genus Pulsadlla they stand between the foliage- 
leaves and floral-leaves. The style of Pulsatilla finally grows out 


ill the form of a feather. The main axis of A. licpatica has unlimited 
growth (it is biaxial), and the flowers are home laterally in the axils of the 
scale-leaves ; in the others (uniaxial) the flower is terminal, and the rhizome 
becomes a syrnpodium after the first flowering Thalictnun (Meadow 

Rue) has no involucre ; 4-5-leaved, greenish perianth. The re- 
ceptacle is flat. The stamens are brightly-coloured and have long filaments ; 
1-5 accessory flowers may occur in the leaf-axils of the panicle-like inflorescence. 

G. CLEMATIDE.E, CLEMATIS GROUP. This differs from all the 
others in the valvate estivation of the calyx and its opposite leaves. 
There are 4 (-several) petaloid sepals ; petals are absent, or 
linear (Atragene). Ovule 1, pendulous. Achenes, often with 
prolonged, feathery style. The majority of the genera are shrubs, 
and climb by their sensitive, twining leaf-stalks. Clematis; 

POLLINATION. The flowers are conspicuous either by coloured petals (honey- 
leaves) (Ranunculus, PtPimia) or coloured sepals (Helleborus, Anemone, Cnltlm, 
etc.), or by both (Aquilerjia, Del2)hiniuin), or by the coloured stamens (Tlialir- 
triini). Some have no honey (Clematis, Anemone, Thalictrum), and are gener- 
ally visited by insects for the sake of their pollen. Others have nectaries 
on the corolla (Ranunculus, TruUiut, Helleborus, Nigella, Aconilnin, etc.), 
more rarely on the stamens (Puls'itilla, Clematis-species), or the carpels 
(Culth'i), or the calyx (certain species of P<ronia). The honey is readily 
accessible in the flat, open flowers, and these flowers also may easily pollinate 
themselves. There is marked protandry where the honey lies deeply hidden, as 
in Aqinli-r/iii, Delphinium, and Aconitum. Helleborus and some Ranunculus- 
species are protogynous. 

About G80 species ; especially in northern temperate climates, and extending 
to the Polar and Alpine regions. Only the Clematitlecc are tropical. 

The order has an abundance of acrid, vesicant properties (R. acer, scclfmtus, 
etc.), and $> lisonous alkaloids (Hell>'borus niger is poisonous). OFFICINAL: 
Aconitum iidpellus (aconitine ; leaves and tuberous roots); the rhizome of 
Hydrciftis canadensis from N. Am. (the alkaloid hydrastine). The order, how- 
ever, is best known for its ornamental plants ; almost all the genera have 
species which are cultivated for their beauty. Sweet-scented flowers are 

Order 2. Nymphaeaceae (Water Lilies). WATER PLANTS; 
generally with large, floating leaves, and large solitary flowers ; 
sepals 3-5, petals 3- GO, stamens 6- GO, carpels 3- GO. The flower 
is hypogynous, but in the Nymphseese different degrees of epigyny 
are found, and from this fact, as well as from the carpels being 
united into one pistil, the family forms a lateral offshoot from 
the Ranunculacete, with much greater modification. The seed 
often has an aril, and, in the majority, a farinaceous nutritive 

W. B. C C 



tissue, partly endosperm, partly perisperm (Fig. 383 C). The 
embryo lias 2 thick cotyledons and a small hypocotyl ; the 
plumule is well developed, with 2-4 leaves. 

1. CABOJIBE.E. 3-4 species (Tropical S. Am.), resembling the Water 
Ranunculus, with two kinds of leaves, the submerged being dissected and the 

rial peltate. The flowers are eucyclic, trimerous, with 2-3 free, epigynous 
carpels. The ovules are situated OH the central line of the carpel an almost 
unique circumstance. Endosperm and perisperm. Ctibomba ; P>rascnia. 

2. NELUMBONEJ:. The leaves are peltate, raised on long stalks 
high above the water. Large, Jiypogynous flowers (Fig. 382) ; 
sepals 4-5 ; petals numerous ; stamens numerous; carpels several, 
distinct. The i^eceptacle is very remarkable, being raised above 
the stamens, and developed into an inverted conical body 011 

the apex of which the nut- 
like fruits are embedded in 
pits. Endosperm is wanting, 
but the embryo is lai'ge and 
has well developed cotyledons. 
Nelunibo, 2 species. N. liitca (N. 
Am.) ; N. speciosa (E. Ind.) was 
sacred amongst the ancient Hindoos 
and Egyptians (the Lotus flower) ; 
its seeds are used as food. 

GROUP. The carpels are united 
into one, many-locular ovary, 
whose numerous ovules are 
situated on the surface of the 
partition walls (as in the Pop- 
pies) ; the stigma is sessile 
and radiating 1 , the number of rays corresponding to the number 
of carpels (Fig. 383). The fruit is a spongy berry with many 
seeds, which have a large perisperm in addition to the endosperm 
(Fig. 383 G). 

Sepals, petals, and stamens often pass gradually over the one 
into the other, the petals becoming narrower by degrees, and 
bearing anthers on each side of the apex, which gradually become 
larger anthers in proportion to the filament, until the perfect 
stamen is developed. The long-stalked leaves are floating, and 
most frequently cordate, elliptical, leathery, with a shiny surface, 
sometimes (as in Victoria regia and Euryale jVrc.c) with strongly 

FIG. 382. Nelumbo nvcifera: vertical section 
through the receptacle. 



projecting thorny ribs on the lower surface. In the intercellular 
passages of the leaves are some peculiar, stellate cells. 

Nuphar has 5 sepals, and an hi/pogynous flower. The petals, which 
are small, have a nectary on the back; the coloured inner side of the sepals 
functions as petals ; the ovate gynoeceum is quite free. N. luteum is a native 
plant (Yellow Water-Lily), with, most frequently, 13 petals and 10-16 locitli in 
the ovary. The rhizome is horizontal, as much as 5-6 cm. in thickness, and 
bears on its under surface a number of roots, which on dying-off leave deep 
scars ; the leaves are borne in spiral lines, and the flowers are solitary in 
certain leaf-axils. The construction of the rhizome is very peculiar ; the 
vascular bundles are scattered and closed as in a monocotyledonous stem. 

A P, C 

FIG. 383. Nympliaia: A flower in longitudinal section, the most external leaves being 
removed ; B fruit ; C seed of Nuphar (longitudinal section) ; the perisperui at the. base, 
the endosperm at the top surrounding the embryo. 

Nymphaea has 4 sepals, and the flower is more or less cpif/y- 
nons. Petals and stamens are inserted at different heights on the 
ovary to just beneath the stigma (Fig. 383). Nymphxa alba (White 
Water-Lily). Victoria regia from the Amazon, and Enryale ferox 
from Asia, have entirely epigynous flowers. The shield-like leaves of 
Victoria areas much as 2 metres in diameter, and the edge is bent up to a 
height of 5-14 cm. ; the flowers are 20-40 cm. in diameter, and change in 
twenty-four hours from white to rose-red. A development of heat, as much as 
14C. above the temperature of the air, together with a strong formation of 
carbonic acid, has been observed during flowering. 

POLLINATION. Nymph<ea alba and other species of the sub-genus Kt/mpJii/- 
topleura are self-pollinated ; the sub-genus Leptopleura is insect-pollinated. 
NitpJutr and Victoria can effect self-fertilisation ; Enryale is self-fertilised, often 
in entirely closed and submerged flowers. The dissemination of the seeds in 
Nnphar luteum is effected by the fruit, which rests on the water, becoming de- 


tacbed from its stalk, and dehiscing from the base upwards so that the seeds are 
set free ; while in Nymph aa aZ&athe spirally-twisted stalk draws the fruit under 
water, and it dehisces by its upper part being thrown off as a hood, and the 
seeds which are enclosed in air-tight, sacs rise to the surface of the water. In 
this condition they are able to float and can only sink to the bottom when the 
air has disappeared. 

53 species ; in fresh water in all parts of the world, but especially in the 
Tropics. The rhizomes and seeds of some may be used as food ; Euryaleferox 
is even cultivated. ]$ympha:a ccerulea and Lotus were sacred among the 

Order 3. Ceratophyllaceae. About 3 species. Aquatic 
plants, submerged, rootless ; leaves cartilaginous, vertieillate, dis- 
sected into repeatedly" dichotomous branches which are finely 
toothed ; only one of the leaves in a whorl supports a vegetative 
branch. The flowers are monoecious, axillary. Inside the 6-12 
perianth-leaves is situated in the g -flower 10-20 stamens with 
thick connective, and in the $ -flower a gynceceum formed by 
one carpel, with one orthotropous and pendulous ovule, which has 
only one integument. Fruit a nut, which, in some species, bears 
on each side a pointed horn, and at the apex a similar one, formed 
by the persistent style. The embryo has an unusually well de- 
veloped plumule with several whorls of leaves. The plant is root- 
less throughout its whole life. CeratnpJiyllum (Horn-wort). 

Order 4. Anonaceae. Sepals 3; petals 3 + 3 (most frequently r air ate) ; 
succeeding these (as in the Ranunciilacca) are numerous acyclic stamens and an 
apocarpous gynceceum ; the flowers are hypogynous, regular and , generally 
very large (2-3 cm. in diameter), and the leaves of the perianth are more or less 
fleshy or leathery. The majority have syncarps with berry-like fruitlets, but 
in Anona and some others the carpels fuse together into a large, head-like fruit 
a kind of composite berry. The seeds have ruminate endosperm as in 
Mi/ristica. Trees or shrubs with alternate, simple, entire, penninerved leaves 
without stipules. 450 (700 ?) species ; especially tropical. The best known are 
Anona clirrimolia, squamosa and reticuJata (all from America) cultivated on 
account of their large, delicious fruits. Some have acrid and aromatic pro- 
perties (Xylnpiti, Cananffi the flowers of the latter yield Ylang-ylang) ; 
Artabotrys odoratissima ; Asimina (N. Am.). 

Order 5. Magnoliacese. Trees or shrubs with scattered, often leathery, 
entire leaves, generally -with stipules, which (as in Ficus) are rolled together and 
form a hood round the 3*0 linger iuternodes above them, and are cast off by the 
unfolding of the next leaf, leaving a ring-like scar. The endosperm is not, 
ruminate. Corolla imbricate. Fruit a syncarp. 

A. MAGNOLIE.E. The flowers are borne singly, and before opening are en- 
veloped in an ochrea-like spathe which corresponds to the stipules of the foliage- 
leaves. The perianth generally consists of 3 trimerous whorls, the external one 
of which is sometimes sepaloid (Liriodcndron, and the majority of Magnolia- 


species), sometimes coloured like the others ; the perianth is sometimes many- 
seriate. Numerous spirally -placed stamens and carpels. The latter are situated 
on the elongated, cylindrical receptacle, and are individually more or less united, 
except in Liriodendroii, where they are free. This last genus has winged achenes ; 
the fruitlets in Magnolia open along the dorsal and ventral sutures, and the 
seeds then hang out, suspended by elastic threads formed from the vascular 
bundles of the funicle and raphe ; they are red and drupaceous, the external 
layer of the shell being fleshy a very rare occurrence. 

B. ILLICIE^; has no stipules. The carpels are situated in a whorl on a short 
receptacle. Follicles, one-seeded. The leaves are dotted by glands containing 
essential oil. Illiciuin ; Drimys. 

70 species; in tropical or temperate climates; none in Europe or Africa. They 
are chiefly used as ornamental plants, e.g. the Tulip-tree (Liriodendroii titlipi- 
fera, N. Am.), Magnolia yrandiftora (N. Am.), Jl/. yulan and fusc.ata (China), 
and others. The remains of Liriodendroii occur as fossils in the Cretaceous 
and Tertiary periods. The fruits of Illiciuin anisatum (Star-aniseed from 
Eastern Asia) are OFFICINAL. The bark of Drimys winteri (S. Am.) is also 
strongly aromatic. 

Order 6. Calycanthaceae. These are very closely related to the Magnoliacese, 
but differ in having perigynous flowers with many perianth-leaves, stamens 
and (about 20) carpels in a continuous spiral, seeds almost devoid of endosperm 
with rolled up, leaf-like cotyledons, and leaves opposite on a square stem. There 
are some species in N. America (Calycanthus Jiorida, occidentalis, etc.) and 
1 in Japan (Chimonaiithus prescox), all strongly aromatic. 

Order 7. Monimiaceae. Aromatic shrubs with opposite leaves. Perigynous 
flowers. The anthers dehisce by valves like those of the Lauracac, and the 
Monimiacese may thus be considered as an apocarpous form of this order. They 
are also closely related to Calycantltacece. 150 species, tropical. Hedycarya, 
Mollinedia, Monimia. 

OrderS. Berberidaceae (Barberries). The regular, $, 
hypogyiious flowers are dimerous or trimerous and have regu- 
larly alternating whorls of free sepals, petals, and stamens and 

FIG. 331. Diagram of Berberis. FIG. 385. Herberts : carpel with 2 stamens. 

1 unilocular carpel : the corolla and stamens have each 2 whorls, 
the calyx at least 2. The anthers open, as in Lauraceae, by 
(2) valves, but are always introrse (Fig. 384). The pistil has 


a large, disc-like, almost sessile stigma (Fig. 385), and in the ovary 
several erect ovules are placed close to the base of the ventral 
suture. The fruit is most frequently a berry. Seeds endosper- 
mous. Shrubs or herbs with scattered, most frequently compound 
leaves (without stipules), and racemose inflorescences. They show a 
relationship to the Lauraceas in the number of the parts of the flower and the 
dehiscence of the anthers. 

Berberis is a shrub ; it has sepals 3 + 3, petals 3 + 3, stamens 3 + 3 
(Fig. 384). The petals (honey-leaves) bear internally at the 
base 2 darkish -yellow nectaries. The filaments are sensitive 
at the base, and suddenly bend inwards if touched at that spot 
(Fig. 385). The racemes often have a terminal, 5-merous flower; they are 
borne on dwarf-branches. The leaves on the long-branches develope into 
thorns, but the buds in their axils, in the same year as themselves, develope as 
the short- branches with simple foliage-leaves, articulated at the base, from 
which fact some authorities have considered that the leaf is compound with a 
single, terminal leaflet Mahonia has imparipinnate leaves. The 
flower has 3 whorls of sepals. Otherwise as in Berberis. 
Epimedium ; herbs with spurred petals ; the flowers dimerous ; 4-5 whorls of 
sepals, 2 of petals and stamens. Fruit a capsule. Leontice, fruit dry. The 
anthers of Podopfiyllum dehisce longitudinally. Nandina. Aceranthits. 

100 species; North temp., especially Asia: fossils in Tertiary. Berberis 
rulgaris is a native of Europe. This and other species, together with Mahonia 
aquifolium (N. Am.), Epimedium alpinum, etc., are cultivated as ornamental 
plants. Several have a yellow colouring matter in the root and stem. 
OFFICINAL: the rhizome of Podophylluni pcltatum (from N. Am.) yields podo- 

Order 9. Menispermaceae. This order has derived its name from the 
more or less crescent-like fruits and seeds. Dioecious. The flowers are 2-3- 
merous, most frequently as in Berberis (S3 + 3, P3 + 3, A3 + 3), with the 
difference that there are 3 free carpels, each with 1 ovule; in some genera, 
however, the number is different. Stamens often united into a bundle (as in 
Myristiea) ; anthers dehiscing longitudinally ; fruit a drupe. The plants (with 
herbaceous or woody stems) belonging to this order are nearly all twining or 
climlimj plants, and have scattered, palmate or peltate, sometimes lobed leaves 
without stipules. Structure of stem anomalous. Cocculus, llenispennum, 
Cissampelus, Anamirta. 

150 species ; Tropical ; very rich in bitter and poisonous properties. OFFI- 
CINAL : Calumba-root from Jateorhiza coluiuba (E. Africa). The following are 
cultivated as ornamental plants : Menispermum canadense (N. Am.) and 31. 
dahitriciini (Asia). The fruits of Anamirta cua-tilus (E. Ind.) are very poisonous 
(" Grains-of-Paradise " ; the poisonous matter is picrotoxine). 

Order 10. Lardizabalaceae. This order, by the free, apocarpous carpels, 
belongs to a more primitive type, and by the united stamens to a more developed 
one. Akebia ; Holbedlia; principally climbing or twining shrubs. About 7 
species in S.E. Asia and S. Am. 



Order 11. Lauraceae (True Laurels). Trees or shrubs; 
the leaves, always without stipules, are simple, most frequently 
scattered, lanceolate or elliptical, entire, penninerved, finely reti- 
culate (except Cinnamomum with 3-5-veined leaf), leathery and 
evergreen (except, e.g. Cinnamomum) ; they are frequently studded 
with clear glands containing volatile oil. The flowers are borne in 
panicles and are small and of a greenish or whitish colour. They 
are regular, perigynous, with most frequently a bowl or cup-shaped 
receptacle (Fig. 386), usually $ , and trimerous (rarely dimerous) 
through all (most frequently 0-7) whorls ; viz. most frequently, 
perianth 2 whorls, stamens 3-4 and carpels 1 (P3 + 3, A3 + 3 + 3 
+ 3, G3) in regular alternation (Fig. 387). Each of the 2 or 4 
loculi of the anthers open by an upwardly directed valve (Fig. 386) ; 
of the stamens, the '2 outermost whorls are generally introrse, 

FIG. 386. Flower of the Cinna- 
mon-tree (Cinnamoinuin zcylani- 
ciuii) (longitudinal section^. 

FIG. 387. Typical dia- 
gram of the Lauraceoe : 

ij staminocles. 

FIG. 388. ini(i-.s 
nobilis : longitudinal 
section of fruit. 

the others extrorse, or 1-3 whorls are developed as staminodes 
(Fig. 3870). The gynoeceum has 1 loculus with 1 style and 1 
pendulous ovule (Fig. 386), and may be considered as formed of 3 
carpels. The fruit is a berry (Fig. 388) or drupe, which often is 
surrounded at its base by the persistent receptacle (as an acorn by 
its cupule), which becomes fleshy and sometimes coloured during 
the ripening of the fruit. The embryo has 2 thick cotyledons, but 
no endosperm (Fig. 388). 

The Lauracejfi present affinities with thePolygonacea?, in which there is found 
perigyny, as well as a similar number of parts in the flower and a similar 
gynceceum, but with erect and orthotropous ovule. From their general charac- 
ters they should be classed among the Polycarpica?, but stand, however, isolated 



by the syncarpous gynoeceum, if it is in reality formed by 3 carpels and not by 1 
only. Hernamliu, which has epigynous monoecious flowers, deviates most. 
(.'axsijtha is a Cuscuta-\\ke, herbaceous, slightly green parasite with twining, 
almost leafless stems. The flower however agrees with the diagram in Fig. 387. 
Some Lauraceffi have curved veins or palminerved and lobed leaves (often 
together with entire ones) c.ij. Sassafras. 

There are 1000 species ; especially iu the forests of tropical S. America and 
Asia, of which they form the principal part. Only Lauras nobilis is found in 
Europe, and there is little doubt that its proper home is in Western Asia. 

FIG. 339. Myristica : fruit. 

FIG. 390. Seed svith aril entire 
and in longitudinal section. 

They are rare in Africa. On account of the volatile oil found in all parts of the 
plant, they are used as spice*, e.g. the false Cinnamon-tree (Dicypellium airyo- 
plii/Uatitni, in the Brazils). The OFFICINAL ones are the Cinnamon-tree (Ciniia- 
momum zeylaiiicum from Ceylon, E. India, Eastern Asia), which is also culti- 
vated ; the Camphor-tree (Ciuuamointtm camphora, Eastern Asia). The Laurel- 
tree (Lauras nohilis, Mediterranean), the berries and leaves of which give 
laurel oil, is medicinal. Scented wood for furniture, etc., is obtained from 
Sassafni* ojlicinalis (from N. Am.). The wood from its roots is officinal. 
Piclmrim "beans" are the large cotyledons of A'cctaudra pichunj, whilst the 


famous " Greenheart " wood of Demarara is the wood of Ncctandru r<>di<ri. 
The pulp and seeds contain a fnttij ml. The pear-like fruit of P<T,~W 
(jratissiina (Mexico, also cultivated) is very delicious. Lindera benzoin is a 
garden shrub ; Lniirns -nobilis likewise. 

Order 12. Myristicaceae (Nutmegs). In this order there is only 1 
genus, niyristica. Trees or shrubs. The leaves agree closely with those of 
the Lauraceae, with which this order has many points in common. The 
majority of the species are aromatic, having in their vegetative parts pellucid 
glands with volatile oils. The flowers are regular, dioecious, trimerous, and 
have a single gamophyllous (cnpular or campanulate) 3-toothed, fleshy perianth. 
Iii the $ -flowers the anthers vary in number (3-15), and they are extrorse 
and borne on a centrally-placed column ; in the ? -flower the gynceceum 
is unilocular, uuicarpellary, with 1 ovule. The FRUIT (Fig. 389) has the form 
of a pear ; it is a fleshy, yellow capsule, which opens along the ventral and 
dorsal sutures, exposing the large seed. This seed has a large, red, irregularly 
branched aril the so-called " mace " ; the "nutmeg," on the other hand, is 
the seed itself with the inner thin portion of the testa, which lias pushed its way 
irregularly into the endosperm, and causes the marbled appearance of the cut 
seed (Fig. 390) ; the external, dark brown, hard, and brittle part of the seed- 
shell is however removed. Mace and nutmeg contain volatile and fatty oils in 
abundance. 80 species. Tropical. The majority are used on account of their 
aromatic seeds and aril, the most important being M. f ray ran s (inoschata), from 
the Moluccas. This is cultivated in special plantations, not only in its native 
home, but in other tropical countries also. Nutmegs were known as com- 
modities in Europe in very ancient times (e.g. by the Romans), but it was not 
until the year 1500 that the tree itself was known. The seed is OFFICINAL. 

Family 10. Rhceadinae. 

The plants belonging 1 to this family are almost exclusively her- 
baceous, with scattered, exstipulate leaves. The flowers are encyclic 
di- or tetra-merous, with the calyx and corolla deciduous, hypoyynous, 
^ ^regular, the gynceceum with 2-several carpels (generally 2, trans- 
versely placed (Figs. 391, 392, 393,397). The ovary is nnilocnlur 
with parietal placenta?, but in Cruciferfe and a few others it becomes 
bilocular by the development of a, false, membranous wall between 
the placentae. The stigmas in the majority of cases are commis- 
sural, i.e. they stand above the placenta?, and not above the dorsal 
line of the carpels. The fruit is nearly always a capsule, which 
opens by the middle portions of the carpels detaching themselves 
as valves, bearing no seed, whilst the placenta? persist as the 
seed-bearing frame. Endosperm is found in Papaveracete and 
Fumariacea>, but is absent in Crucifene and Capparidacex. This 
family through the Papaveraceae is related to the Polycarpica? (the Nymphseacese), 
through the Capparidacese to the Resedacere in the next family. 

Exceptions to the above are: Escltscholtzia, Subuhiriu (Fig. 403) and a few 



CapparidaoesB, in which perigynous flowers are found. A f^w Papaveraceae and 
Fumariaceje have trimerous flowers. In Fumaria and certain Cruciferae, the 
fruit is a nut. The Fumariaceae have zygomorphic flowers. Trees and shrubs 
are almost entirely confined to the Capparidacese, in which order stipules also 
are found. 

Order 1. Papaveraceae (Poppies). Herbaceous plants with 

stiff hairs and latex ; flowers regular (Fig. 391) with generally 2 
(-3) sepals (which fall off as the flower opens), "2 + 2 petals (im- 
bricate and crumpled in the bud) ivitliout spur, numerous stamens in 
several alternating whorls (generally a multiple of 2) ; carpels 2- 
several, united into a unilocular gynceceum. Trimerous flowers 
also occur. Capsule with very numerous seeds on the parietal 
placentae ; embryo small, with large, oleaginous endosperm (Fig. 
392). The leaves have no stipules and are generally pinnately 


FIG. 391. ,4 Diagram of the flower of 
Glaucium and the clichasium (which 
becomes transformed into a scorpioid 
cyme). -B Pupa re f ai'ijcmone, transverse 
section of the ovary with indication of 
the position of the stigmas. 

FIG. 39.'. Papaver somniferum : A cap- 
sule ; st the stigma; o> valves; h pores ; 
-B seed in longitudinal section ; a!b endo- 
sperm; emb embryo. 

Papaver (Poppy) has large, solitary, terminal flowers ; petals 
firmly and irregularly folded in aestivation ; gynceceum formed by 
many (4-15) carpels ; stigmas velvety, sessile and stellate (the rays 
stand above the placenta) (Fig. 391 U). The edges of the carpels 
project deeply into the ovary, but do not meet in the centre, so 
that it remains unilocular. The capsule opens by pores placed 
close beneath the stigma, and formed of small valves alternating 
with the placentae and the rays of the stigma (Fig. 391). P. dubium, 
P. anjemone, P. rhceas. Chelidonium (Greater Celandine) has 
yellow latex, flowers in umbellate cymes (the terminal, central flower 
opening first) and only 2 carpels ; the fruit resembles the siliqua 

R1KEADINJ!. 395 

of the Crucifera? in having two barren values, which are detached 
from the base upwards, and a seed-bearing frame, but there i.s no 
partition wall formed between the placentae. Ch. majus. The 
majority of the other genera have, like Chelidoniti-ni. 2 carpels (lateral and alter- 
nating with the sepals: Fig. 391 A) and siliqua-like fruit, thus : Escltselioltsia 
(perigynous) with a linear, stigma-bearing prolongation extending as far above 
the placenta? as above the dorsal suture of the carpels; Glaucium (Horn-Poppy) ; 
G. lutenm, whose extremely long, thin capsule differs from that of Chelidonium 
by the formation, during ripening, of a thick, spongy (false) replum, which 
persists when the valves are detached ; Sanguinaria with red latex, the 2 petals 
divided into 8-12 small petals (perhaps by dedoublement) ; Miicleya and 
Bocconia (1-seeded capsule) with 2 sepals and no petals. Trimerous flowers 
are found in Argemone and Platystemon (with a curious fruit, carpels free, and 
transversely divided and constricted into joints which separate as nut-like Tpot- 
lionsj.Meconopsia.Hypecoum (Fig 393 C) has tri-lobed and three cleft petals, 
4 free stamens with 4-locular anthers and a jointed siliqua; it presents a tran- 
sitional form to the Fumariaceas, with which order it is sometimes included. 

POLLINATION. Papavcr and ClicJidoninm have no honey, and are without 
doubt only visited by insects for the sake of the pollen. The anthers and 
stigmas mature about the same time. There are 80 species ; especially from 
warm climates. OFFICINAL : Papaver soninifemm (Opium-Poppy) ; the latex 
of its unripe capsules is obtained by incisions, and dried (opium) ; it contains 
many alkaloids : morphine, papaverine, narcotine, thebaine.etc. The oleaginous 
seeds are also used in the manufacture of oil. Its home is in the East, where 
it is extensively cultivated. The petals of the Corn-poppy (P. rlia'as] are also 
officinal. Several species are cultivated as ornamental plants. 

Order 2. Fumariacse (Fumitories). This order differs from 
the closely allied Papaveraceae in the absence of latex, a poorer 
flower, generally transversely zygomorphic (Fig. 393 Z/), in which 
case one or both of the outer lateral petals are gibbous, or pro- 
longed into a spur; the stamens are especially anomalous. Sepals 
2, caducous ; petals 2 + 2 ; stamens 2, tripartite ; each lateral anther 
is bilocular (Figs. 393 A, B ; 395) ; gynoeceum bicarpellate. 
The fruit is a nut or siliqua-like capsule. Endosperm. Herbs with 
scattered, repeatedly pinnately-divided leaves without stipules, 
generally quite glabrous and glaucous ; the flowers are arranged 
in racemes with subtending bracts, but the bracteoles are some- 
times suppressed. 

Diceutra (syn. Dielytra) and Adliimia have a doubly symmet- 
rical flower, with a spur or gibbous swelling at the base of 
each of the laterally-placed petals (Figs. 393 A, 394). Cori/dalis 
has a zygomorphic flower, only one of the lateral petals having a 
spur, and consequently there is only one nectary at the base of the 
bundle of stamens, which stands right in front of the spur (Fig. 



893 .B, 39."), 396). The fruit is a many-seeded siliqua-like capsule. 
A peculiarity of the flower is that the plane of symmetry passes transrerseli/ 
through the flowers, whilst iu nearly all other zygomorphic flowers it lies in 
the median line. Moreover, the flower is turned, so that the plane of symmetry 
ultimately becomes nearly vertical, and the spur is directed backwards. Many 
species have subterranean tubers ; in these the embryo germinates with one 

FIG. 393. Diagram of Dicmfra (A), Corydalis (B), and HyjuTOitm (C). 

cotyledon, which is lanceolate and resembles a foliage-leaf. The tuber is in 
some the swollen hypocotyl (C. cara), in others a swollen root (C. fabncea, 
etc.), which grows down through the precisely similar swollen root of the 
mother-plant. The sub-genus Ceratocapnos has dimorphic fruits (nuts and 
capsules) in the same raceme. Fumaria differs from Corydalis only by 
its almost drupaceous, one-seeded nut (Fig. 395). 

THE STRUCTURE OF THE FLOWER. Hijpecoum among the Papaveraceas is the 
connecting link with the Funiariacerc. The diagram (Fig. 393 C) corresponds 
both in number and in the relative position of its members with that of most 
of the other Papaveraceas (Fig. 391), except that there are only four stamens 
(with extrorse anthers). In Dicentrn (Fig. 393 A), the two central (uppermost) 
stamens are absent, but each of the two lateral ones are divided into three 
filaments, of which the central one bears a four-locular anther, and each of the 

FIG. 394. }>icent,-a speeMiilii : A flosver (f ) ; B the same, after removal of half of one 
outer petal; the cap, formed by the inner petals, is moved away from the anthers and 
stigma; the insect does this with the lower side of its abdomen, aud thus rubs the stigma 
on the hairs of its ventral surface; the dotted line at e indicates the direction of the 
proboscis; C andrceciuin andgynoeceum ; D stigma. 



others a two-locular (half) anther. Cori/dalis and Fumaria stand alone in the 
symmetry of the flower, differing from Dicentnt in having only one of the lateral 
petals (Fig. 893 B, sp) prolonged into a spur, while in Dicentra both the 
petals are spurred. This structure has been interpreted in various ways. 
According to Asa Gray the median stamens are absent in the last-named 
genera, and the lateral cues are split in a similar manner to the petals of 
Hypecomn. Another, and no doubt the most reasonable theory (adduced by 
De Candolle), is : that two median stamens are split, the two parts move 
laterally, each to their respective sides and become united with the two lateral 
stamens ; this affords a natural explanation of the two half-anthers, and estab- 
lishes a close relationship to the Cruciferas. A third interpretation, held by 
Eichler and others, is as follows : the median stamens are always wanting ; 
when they appear to be present, as in Hypecouni, it is due to the fact that the 

FIG. 393. .Fumaria officin- 
alis : A the flower in longi- 
tudinal section ; B the anrlroe- 
cium nnd gvnoeceuin ; nectary 
to the right. 

FIG. 396. Conjdalis cava: a a flower (Literal viaxv); 
b the anthers lying round the stigma ; o the anthers 
shortly before the opening of the flower; A the head 
of the stigma; p relative position of the parts of the 
flower during the visit of an insect. 

side portions of the lateral stameus approach each other (as interpetiolar 
stipules) and coalesce into an apparently single stamen. 

130 species ; mostly from the northern temperatures. 

POLLINATION. Fumaria, with its inconspicuous flowers, has to a great 
extent to resort to self-pollination. Cori/dalis, on the other hand, is dependent 
oil cross-pollination ; C. cava is even absolutely sterile with its own pollen. 
Corydalis is pollinated by insects with long probosces (humble-bees, bees), 
which are able to reach the honey secreted in the spur ; as they alight on the 
flowers they press the exterior petals on one side (Fig. 396 e}, so that the 
stigma, surrounded by the anthers, projects forward ; the proboscis is intro- 
duced in the direction of the arrow in the figure, and during this act the 
under-surface of the insect is covered with pollen, which is transferred by similar 


movements to the stigma of another (older) rower. Ornamental plants ; 
Dicentra (^ectabilis and eximia), Adlumia, Corydalis. 

Order 3. Cruciferae (Crucifers). The flowers are regular, 
5; sepals 4, free ('2 + 2), deciduous; petals 4, free, deciduous, 
unguiculate, placed diagonally in one whorl, and alternating 1 with 
the sepals ; stamens 6 ; the 2 outer are short, the 4 inner (in reality 
the two median split to the base) longer, placed in pairs (tetrad}' - 
namia of Linnaeus) ; gynceceum syncarpous formed by 2 (as in the 
previous order, lateral) carpels, with 2 parietal placenta?, but 
divided into two loculi by a spurious membranous dissepiment 
(replum) (Fig. 397). Style single, with a capitate, usually two- 
lobed stigma, generally cornmisural, that is, placed above the 
parietal placenta? (Fig. 397), but it may also be placed above the 
dorsal suture, or remain undivided. Ovules curved. The fruit is 
generally a bivalvular sill qua (Fig. 398 B, C), the valves separat- 
ing from below upwards, and leaving the 
placentae attached to the replnm ; other 
forms of fruits are described below. The 
oily seeds have no endosperm (endosperm is 
present in the two previous orders) ; the 
embryo is curved (Figs. 398 E, F- 399, 400). 
In general they are herbaceous plants, 
without latex, with scattered, penninerved 
leaves, without stipules ; the inflorescence 
FIG. 39/.-Diagram of a ig y characteristic, namely, a raceme 

Cruciferous flower. 

with the flowers aggregated together at the 

time of flowering into a corymb, and destitute of both bracts and 

Many are biennial, forming in the first year a close leaf-rosette. By culti- 
vation the tap-root can readily be induced to swell out into the form of a tuber 
(Turnips, Swedes, etc.). Sti})iiles are found indicated by small glands on the 
very young leaves ; in Cochlearia armoracia they are fairly large triangular 
scales. Stellate hairs often occur. Floral-leaves are occasionally developed. 
Terminal flowers are never found in the inflorescences. Iberis and Teesdalia 
have zijgomorphic flowers. Subulada (Fig. 403) is perigynous. The 2 external 
sepals (Fig. 397) stand in the median plane ; it may therefore be supposed that 
there are two bracteoles outside these which, however, are suppressed, and can 
only in a few instances be traced in the young flower ; the two lateral sepals 
are often gibbous at the base, and serve as reservoirs for the nectar secreted 
by the glands placed above them ; they correspond in position to the external 
petals of the Fumariacefe. The 4 petals which follow next arise simultaneously, 
and alternate with the 4 sepals ; if it could be shown that these are merely 2 
median petals, which have been deeply cleft and the two parts separated from 


each other and displaced to the diagonal position, there would be a perfect 
correspondence with the Fumariaceous flower ; then the petals would be followed 

FIG. 398. Bfassica oleracea -. A raceme ; B, C siliqua ; D seed ; E embryo ; F transverse 

section of seed. 

in regular alternation by the 2 lateral small stamens, the 2 median long stamens, 
which it has been proved are split into 4 and placed in couples, and the 2 
laterally-placed carpels, in all 6 dimerous whorls. But the formation of the 

Fi&. 400. Transverse section of seed of 

FIG. 399. Transverse section of seed and 
embryo of Cheiranthus cheiri. 

corolla by the splitting of 2 petals does not agree with the development of the 
flower or bear comparison, and hence the only fact in favour of this theory is 


the otherwise prevailing correspondence with the Fumariacese. Yet it may be 
observed that in special cases eacli pair of long stamens clearly enough arises 
from one protuberance and even later on may be considerably united or entirely 
undivided (<'.</. Telia) ; in other instances tbey are quite distinct from the 
beginning, and it is possible that this latter condition has become constant in 
the corolla. Lepidium ruderale and others have no corolla. Senebiera didijma 
has only 2 median stamens. Megacarpcea has several stamens, no doubt by 
drdoublement, as in Capparidacese. The number of carpels may also be 
abnormally increased ; Tetrupoma larltarei folium has normally 4 carpels with 
an equal number of placenta? and repla. It is supposed to be a variety of 
Nasturtium palitstrc. The 2-4-8-10 greenish gland*, which are found at the 
base of the stamens, are nectaries, morphologically emergences, and not rudi- 
mentary stamens. The forms of fruit* are of great systematic significance, see 
the genera. In some species dimorphic fruits are present, e.g. Cunlaiuhie 
c'lennpodiifolia which has both ordinary Cardamine-BiMqu&B and 1-seeded 

The curved embryo appears in five forms, which have systematic importance : 
1. To the PLEDRORHIZJE belong those genera whose radicle (with the hypocotyl) 
lies bent upwards along the edge of the fiat cotyledons (Fig. 399) ; to this group 
belong Cardamine, Nasturtium, Oheiranthus, JIatthiola, Cochlearia, Draba, 
Iberis, Thlaspi, etc. ; diagrammatic transverse section : Q 2. To XOTO- 
EHIZ.B belong those whose radicle lies in an upward direction along the back of 
one of the flat cotyledons (Figs. 400, 413) ; e.g. Hesptri*, Sisymbrium, Lepidium, 
Capsella, Camelina : Q !! 3. ORTHOPLOCE.E differ from the Notorhizere in 
having the cotyledons folded (not flat) (Fig. 398 E, F) ; to this belong Hrai- 
xica, Simipix, lidplmnus, Crainbc, etc.: Q^- 4. SPIROLOBE.E : the radicle lies 
as in the Notorhizte, but the cotyledons are so rolled together that a transverse 
section of the seed cuts them twice ; Bunias : Q I! II- 5. DIPLECOLOBE.E : the 
cotyledons are folded forward and backward so that a transverse section cuts 
them several times; Subularia, Svnebitm : Q |] || ||. 

On germination the cotyledons appear above the ground as 
green leaves ; in the Orthoploceae they are bilobed, in the Lepldinin- 
species divided. 

1. Silicula, broad replum (Siliculosse'latiseptse),. valves flat 
or slightly vaulted, and the replum extends through the greatest 
width of the silicula (Fig. 404). The seeds are situated in two rows. 

O ~~ ' Cochlearia (Horse-radish) : the siliqua is nearly spheroid ; 
glabrous herbs, generally with flesh}-, stalked leaves, and white 
flowers. Draba has an oblong, lanceolate, somewhat compressed 
silicula ; herbs with small rosettes of leaves, most frequently with 
stellate and long-stalked racemes. Alyssum and Berteroa are 
whitish, on account of the stellate hairs ; they have a more com- 
pressed and round or elliptical silicula. Vesicaria ; Anbrietin. 
Lunaria (Honesty, Fig. 401) : very broad and flat silicula with 
long stalk (the receptacle as in Capparidacese). 



O |[ : Camelina (Gold -of -pleasure) has a spheroid, pear-shaped 
siliqua with a small rim passing right round (Fig. 402). Subu- 
laria (Awlvvort), an aquatic plant with perigynous flower (Figi 
403) and folded cotyledons. 

2. Silicula, narrow replum (Siliculosee angustiseptse), i.e. 
the replum is much shorter than the arched, more or less boat- 
shaped valves (Figs. 405, 406, 407). 

O=: Thlaspi (Penny-Cress) has a flat, almost circular silicula, 
emarginate or cordate, with a well-developed wing round the edge 
(Fig. 406). Iberis and Teesdalia : the racemes during flowering 

PIG. 401. Lunaria biennis. 
Fruit, the valves of which 
have fallen off. 

FIG. 402. Came- 
lina sativa. Fruit. 

FIG. 4fl3. Sulndaria aquatica. 
Longitudiual section through 
the flower. 

are especially corymbose, and the most external petals of the outer 
flowers project radially and are much larger than the other two 
(the flower is zygomorphic). Biscutella, Megacarpcea. 

O || : Capsella (Shepherd's-Purse) has a wingless, obcordate or 
triangular silicula (Fig. 407). Lepidium (Pepperwort) has a few- 
(2-4) seeded, slightly winged, oval silicula. Senebiera has a silicula 
splitting longitudinally into two nut-like portions ; its cotyledons 
are folded. Anastatica hierochuntica ("Eose of Jericho ") is au annual, sili- 
cula-fruited, desert plant (Arabia, Syria, N. Africa). After the flowering all its 
then leafless branches bend together upwards, forming a kind of ball; this 
spreads out again on coming in contact with water, and the fruits then dis- 
seminate their seeds, which germinate very quickly, often in the fruit. 

W. B. D D 



3. Siliqua (Siliquoso?). The fruit is a true siliqua, several 
times longer than broad. The seeds in most are borne apparently 
in one row. 

O : Brassica (Cabbage). The seeds are placed apparently 
in one row in each loculus (Fig. 398 G) ; the style is long and round ; 
the valves have only 1 strong, longitudinal rib. Melanosinapis (If. 



FIG. 401. Transverse section of a silicula 
with broad replum : s replum ; k the valves. 

FIG. 405. Transverse section of a 
silicula with narrow replum. 

nigra, Black-mustard); the style is compressed, two-edged; the 
valves of the siliqua are one ribbed. -Sinapis (Mustard) ; quad- 
rangular or flat style (in which in most cases there is a seed) and 
3-5 strong, longitudinal ribs on the valves. Eruca differs from 
Brassica by the shorter siliqua, broad, sword-like " beak " and 
seeds in two rows. 

O=- (Fig. 399) : Cardamine (Bitter Cress) has a long, linear 
siliqua, with flat, unribbed, elastic valves. The leaves are most 
frequently pinnatifid or pinnate. C. 2 yratensis reproduces by buds 
formed in the axils of tbe leaves. Arabis (Rock Cress); Mattliiola 
(Stock); Cheiranthus cheiri (Wallflower); Barbarea (Winter Cress) 

FIG. 406. Tlilaspi anenfe. 

FIG. 407. Silicula of Capsella 

(double-edged, quadrangular siliqua) ; Nasturtium (N. officinal e, 
Water-cress) ; the siliqua of the latter genus is in some species 
short, in others long. 

O || (400) : Sisymbriuni (Hedge Mustard) the valves of the 
siliqua are 3-ribbed. Erysimum ; llesperis ; Schizopetalum (with 
iimbriate petals). 


4. Fruit jointed (Lomentaceae). The fruit is divided by 
transverse walls into as many spaces as there are seeds, and de- 
hisces at maturity, generally transversely, into a corresponding 
number of nut-like joints (" articulate-siliqua.'') 

O : Crambe (Kale, Fig. 408). The fruit has only 2 joints. 
The lower one resembles a short, thick stalk, and is barren, the 
upper one is spherical, and has 1 seed. Cakile (fl. maritime/,, Sea- 
kale) ; the lower node is triangular, 1-locular, the upper one more 
ensiform, 1-locular (Fig. 409). 

O : R'tphanus has a long siliqua, which, in the garden 
Radish (E. sativus), is spongy and slightly abstracted (Fig. 411), but 

FIG. 408. Fruit of 
Crambe maritima. 

FIG. 409. Caltile FIG. 410. Raphanus FIG. 411. Eaphanus 
maritima. Fruit (f). raphanistrum. sativus. 

neither opens nor divides transversely (a kind of dry berry), and 
which in the Wild Radish (R. rapkanistrum) (Fig. 410) is abstricted 
in the form of a string of pearls, and separates into many joints. 
I?, sativus ; the " Radish " is formed by the hypocotyl, after the bursting of 
its external, cortical portions (of which there are generally two patches at the 
top of the Radish). 

5. Siliqua indehiscent (Nucumentacere). The fruit is a 
short, unjointed, unilocular and 1-seeded nut, and the fruit-stalks 
are often long, slender, and drooping. (Sometimes a thin endo- 
sperm is present). Isatis (Woad) has most frequently an oblong, 
small-winged nut; O || (Figs. 412, 413). Bunlas ; Neslia. 



[The systematic division of this order given above is founded upon that of 
A. P. de Candolle. Prantl (En pier and Prantl, Nat. Fain.), 1891, adopts a some- 
what different system, which may briefly be summarised as follows : 

A. Hairs uubranched or absent; no glandular hairs. 

1. TIIELYPODIE.E. Stigma equally developed on all sides ; style undivided 

or prolonged above the middle of the carpels, or turned back. 
Stanleyiiice ; Heliophilinee. 

2. SINAPE^E. Stigrua strongly developed above the placenta ; style beaked 

or two-lobed. 

a. Cotyledons arising behind the bend of the embryo. Lepidiince. 
I. Cotyledons arising at the bend of the embryo. 

a. Only lateral nectaries. Generally a silicula or iudehiscent 

fruit. Coclileariinif. 

13. Generally a siliqua, more rarely a silicula or transversely- 
divided or indehiscent fruit. Nectaries generally lateral and 
median. Alliariimc ; Sixymbriinic ; Velllmc; Brusslcince ; 

11. Hairs collectively or partially branched, very rarely entirely absent ; 
glandular hairs are sometimes also present. 


2. HESPEBIDE^. Stigma strongly developed above the placenta ; style un- 

divided or prolonged above the placentae into shorter or longer lobes. 
a. Surface cells of the replum, not divided diagonally. CapselUnte ; 

Tarritlnce; EryximiiiiC ; Alysxhuc. 
1>. Surface cells of the replum divided diagonally. Malcolmiinee ; 

Helper idirtcc ; Moricaiidiiiue.] 

POLLINATION. Honey is secreted by the nectaries mentioned above; but the 
position of the stamens is not always the most fa< curable for pollination by 

insects (in these flowers the honey-seeking 
insect must touch tl;e anthers with one of its 
sides and the stigma with the other), and 
self-fertilisation is common. In some species 
(Cardamine praiensis] the long stamens turn 
their anthers outwards towards the small 
stamens, so that 3 anthers surround each of 
the two large entrances to the nectaries. 

1'200 species (180 genera), especially in 
the cold and temperate parts of the Old 
'World (Europe, W. Asia). Many are weeds 
in this country, e.g. Wild Cabbage (Urtuxii n 
cnmpesiris), Charlock (Sinajiis <, Wild 
Eadish (RapJiamis rapJianiiitrum) and others. 
The order is acrid and oleaginous. Oil is 
obtained from many of the oil-containing 
seeds, especially of the Rape (Brassica napus), 
Summer-Bape (the oil-yielding cultivated form 
of the Field-Cabbage) and Camelina. Several 
are pot-herbs or fodder plants, e.g. Cabbage 

U2. 413. 

FIG. 412. Isntis tiiirlnrin. Fruit 
(Kitr. 412) ; and in longitudinal sec- 
tion Fig. 413). (Mag.) 



(Brasxica olcracea) with its numerous varieties : Cauliflower (var. botrytis ; the 
entire inflorescence is abnormally branched and fleshy), Kohlrabi (var. yonyij- 
lodes, with swollen, tuberous stem), Kale, Red-Cabbage, White-Cabbage, etc. ; 
H. campestrin, var. rapifera (Turnip) ; 11. napus, var. rapifera (Swede) ; Eapha- 
nus satious (Radish from W. Asia), 11. caudatus (long Radish) ; Nasturtium 
officinale (Water Cress), Lcpidium sativum (Garden Cress), and Barbarea prcecox 
(Early Cress) ; Crambe muritinia (Sea Kale). The seeds of the following are 
especially used as spices : (the flour of) Melanosinapis (Black-mustard), and 
Si mi pis alba (White-mustard), which are officinal like the root of Cochlcaria 
armoracia (Horse-radish, E. Eur). The herbaceous parts of Cochlearia officiiiiilis 
and dniiica are medicinal. A blue dye (woad) is extracted from Isatis. Orna- 
mental plants: Clieiranthns clieiri (Wallflower), Mutthiola (Stock), Iberig, 
Hesperi*, Limaria, and others (especially from S. Eur.). Sweet-scented flowers 
are rave. 

FIG. 414. Gynandropsis FIG. 415. Capparis spinosa. 


Order 4. Capparidaceae (Capers). The relationship with the Cruciferaj 
is so close that certain forms are with difficulty distinguished from them. The 
diagram of the flower is the same in the number and position of its parts, but 
it differs in the modifications which occur in the development of the stamens. 
In some genera all 4 stamens are undivided ; in others both the 2 median ones 
are divided as in the Cruciferaa (6 stamens, but not tetradynamous) (Fig. 414) ; 
in other genera only 1 of these ; in other instances again they are divided into 
more than 2 ; and finally the 2 lateral ones also may be found divided, so that 
indefinite stamens occur (Fig. 415). The bi-carpellate gyuoaceum is unll ocular 
(without replum), but more than 2 carpels may occur. The ovary is elevated 


on a stalk (sometimes as much as 1 foot in length) ; also between the stamens 
and corolla a similar stalk may be found (Fig. 414). The fruit is long and 
siliquose (Cleome, Polanma, Gynandropsis), or a berry (Capfaris). Endos- 
perm absent. Some have zygomorphic flowers. Gamosepalous calyx and 
perigynous flowers also occur. 350 species ; especially in the Tropics. The 
majority are trees and shrubs, and they differ also from the Cruciferse in having 
distinct stipules present in some species. 

" Capers " are the flower buds of the climbing, thorny shrub, Cctpparis spinosa 
(Fig. 415), which grows in the Mediterranean. 

Family 11. Cistiflorae. 

The flowers in this family are perfect, regular (except Resedacew, 
Violacete), hypogynons, the perianth-leaves free (a few have them 
slightly united), {estivation most frequently imbricate ; they are 
eucyclic in the androecium, and most frequently in the other parts, 
and generally 5-merous with So, P5, A 5 + 5, Gr3, but other numbers 
also occur ; several have indefinite stamens, but the stamens ai'ise 
(where the development is known) in centrifugal order and are 
arranged, often very distinctly, in bundles ; in other words, the 
large number of stamens is formed by the splitting of a small 
number (most frequently 5) ; a true spiral arrangement is never 
found. Grynceceum syncarpous, multicarpellary (Dilleniacete and 
a few Resedaceie are apocarpous), most frequently the number of 
carpels is 3, forming a unilocnlar ovary with parietal placenta 1 , 
but parallel with this, multilocular ovaries, with the ovules placed 
in the inner angle of the loculi, are also found, and a few genera 
have a free, centrally-placed placenta. The fruit is most frequently 
a capsule. The dehiscence is never with a " replum," i.e. the 
persistent frame of the placenta, as in the family Rhoeadinte. One 
half of the orders has endosperm (Violacew, Cistacew, Droseracefe, 
Bixacete, Ternstroemiacea?, etc.), the other has no endosperm 
(Resedacefe, Hyper icacete, Elatinaceog, Tamaricacea>, etc.) ; some 
have a curved, the majority a straight embryo. The family is 
scarcely quite natural ; in the future the orders will probably be arranged 

Order 1. Resedaceae (Mignonettes). -- Herbs or small 
shrubs with spirally-placed leaves and very small, gland-like 
stipules (as in Cruciferae) ; the $ , hypogynous flowers are sygo- 
morphic, and arranged in racemes or spikes t} T pically without 
bracteoles. The zygomorphic structure is produced by the greater 
development of the posterior side of the flower, especially the petals 
and the nectary (" disc," in Fig. 416 d) which is situated between 


the petals and stamens ; in general there are 5-8 free sepals 
and petals, the latter consisting of a large scale-like sheath with 
a fimbriated blade (see Fig.) ; stamens numer- 
ous ; carpels 6-2 united together; ovary uni- 
locular with parietal placentae, but the cavity 
of the ovary is not closed at the top. In Astro- 
carpus the gynoeceum is apocarpous. The fruit 
is most frequently a capsule ; the seeds are 
reniform, without endosperm, and the embryo 
is curved. 

This order connects the Bhceadinns with the Cisti- 

FIG. 41<5. Diagram of 
rlora}. It is closely allied to the Rhceadmas by its Reseda odorata. 

external appearance, even by the smell and taste, the 

parietal placeutation, structure of the seeds, the inflorescences, etc., whilst by 
the irregular flowers and the disc placed at the posterior side of the flower, 
it is cllied to Cappuridacete, but differs from this order in not having its 
characteristic number (2-4) and by the very different mode of dehiscence of 
the fruit, etc. It differs from the other orders of this family chiefly in the 
fact that the number of the perianth-leaves is not constantly 5. In R>seda 
luteol/t both the calyx and corolla appear to be 4- leaved, because the posterior 
sepal is suppressed, and the 2 posterior petals are united. Where tin re are 10 
stamens, they stand in 2 whorls, i.e. in front of the sepals and petals ; if there 
are several, their position depends upon the splitting. Astrocarpus is remark- 
able for its apocarpous fruit and the position of the ovules on the dorsal suture 
of the carpel. 

The yellow, flat disc at the back of the flower serves as a nectary, the honey 
being protected by the lobes of the petals. If pollination by insects is not 
effected, then self-pollination may take place, at all events in R. odomta. 

45 species ; the majority in the Mediterranean and in Persia. Reseda odnratu 
(from Egypt) is cultivated on account of its sweet scent; R. lutcola ("Dyer's 
Weed ") yields a yellow dye. 

Order 2. Droseraceae (Sundews). Herbs, chiefly living on 
moors or in water, and whose leaves are adapted to catch and 
digest small animals. With regard to the flower, they are closely 
allied to the Violacete, especially to those with regular flowers. 
Drosera (Sundew) has a long-stalked scorpioid cyme with regular, 
^ , hypogynous flowers, 5-merons as in Viola. S5, Po, A5, G3 
(in a syncarpous gynosceum, with free, bifid styles and basal or 
parietally-placed ovules in the unilocular ovaries). The capsule 
opens also as in Viola, but, among other differences, the styles are 
free, the seeds very small, and surrounded by a loosely lying, thin 
shell. Droser.i has radical, long-stalked leaves with the blade (Fig. 417) 
covered by numerous strong glandular hairs, placed on the edge and in the 
middle ; when small animals are caught by these hairs, the latter and the entire 



blade close slowly over them dissolving and absorbing all the digestible matter 
as nourishment. 

Dioncea muscipula (Fly-trap ; N. Am.) has the same appearance as Drosera, 
but the leaves are constructed as in Fig. 418. The stalk is flat and winged, the 
blade small, circular, with powerful, pointed teeth along the edge, and on its 

FIG. 417. Leaf-rosette of Drosera rotundifolia (nat. size), and a leaf (magnified). 

surface are 6 small bristles (A), which are very sensitive. When these are 
touched the blade quickly closes, folding along the midrib (B, C) and im- 
prisoning the irritating object, the teeth round the edges fitting like the teeth 
of a trap. If it happens to be an insect or similar body, a digestive fluid is 
secreted which, like the gastric juice, dissolves the digestible portions. Aldro- 
r/i/uUa I'esiculosa (Central and S. Europe) captures small aquatic animals 
in a similar manner ; it is a floating, aquatic plant, the two halves of its leaves 
also close together when irritated (Fig. 419). Drosophyllum. 
About 110 species ; most of them in the temperate regions. 

FIG. 418. Dloncea muscipula. Leaves (nat. size). 

Orders 3 and 4. Sarraceniaceae and Nepenthaceae. These two orders 
are perhaps most closely allied to the Droseracese and agree with these, among 
other things, in the manner of taking nourishment. Like the Droseracea? they 
absorb nitrogenous food from dissolved animal matter by means of their leaves, 
which are specially constructed both to catch, to retain, and to digest any small 



animals which may be caught. The SARRACENIACE.E are North American 
marsh-plants (10 species) which have pitcher-like leaf -stalks, in the cavity of 
which a fluid (with properties approaching those of gastric juice) is secreted, 

FIG. 419. Aldrovand ; a vesiculosa : A a plant (nat. size). JSLeaf 
(mag.); the blade is closed; the winged stalk is prolonged into 
4-6 irritable bristles. 

and which bear at the apex a small, lid-like blade ; these leaf-stalks are the 
catching and digestive organs. Sarracenia, Darlingtonia. 

FIG. 420. .Nepenthes (reduced). 

NEPENTHACE^; has only 1 genus, Nepenthes (the Pitcher-plant ; about 35 
species), especially found in tropical E. Asia; the majority are climbing shrubs. 
The leaf- stalks are twining organs, and terminate either simply in a tendril, 
or in addition to this, with a pitcher- shaped body (which in some species maybe 



as much as a foot in length) on whose upper edge a lid-like structure is found 
(Fig. 420). In this pitcher, as among the Sarraceuiaceas, a fluid is secreted 
which is able to digest the animals captured (sometimes rather large) and which 
corresponds in some degree to the gastric juice. 

Order 5. Violacese (Violets). The flowers are ^ > an d gener- 
ally zygomorphic, hypogynous, with S5, P5, A5, G3 (Fig. 421). 
The stamens are closely applied to the ovary, they have a very 
short filament, and at their summit generally a membranous 
appendage formed by the prolongation of the connective (Fig. 
422 g). The ovary is unilocular with 3 parietal placenta? ; style 

FIG. 421. Diagram of 

FIG. 422. The large-flowered form of Viola tricolor : A the 
flower in median longitudinal section ; B the gynoeceum. 

undivided (Fig. 422 B). The fruit is usually a 3-valved capsule, 
opening along the dorsal sutures (Fig. 423). Embryo straight ; 
endosperm fleshy (Fig. 425). Many are herbaceous plants (e.g. 
Viola), but in the Tropics shrubs are also found (e.g. lonidium) ; 
a few are lianes ; the leaves are scattered, with stipules, and 
involute in the bud. 

Viola. The sepals are prolonged backwards beyond the point of 
insertion (appendiculate) ; the corolla is polypetalous, descending 
imbricate, and zygomorphic, its anterior petal being larger than the 
others and provided with a spur (Fig. 421). The 2 anterior of the 
5 almost sessile stamens are provided with a spur-like nectary, 
which protrudes a considerable distance into the petaloid spur 
(Figs. 421, 422 n, sp). The style is club-like, and bears the 



stigma in a groove on the anterior side (Fig. 422 st). Herbs 
with rhizomes, or annuals; flowers solitary. V. odorata, caiiina, etc., have 
cleistogamic flowers which produce fruit in addition to the large, coloured 
(violet) flowers. The Pansy (V. tricolor) has large flowers adapted for insect- 
pollination, and also smaller, less conspicuous ones designed for self-pollina- 
tion. The stigma, as in Fig. 422 A, st, and B, is situated on the anterior 
side of the stylar-head, immediately in front of the channel leading down to 
the spur (sp) ; below it is situated a valve, easily covered with pollen when the 
proboscis of an insect is introduced into the spur, but which closes upon its 
withdrawal ; cross-pollination is thus secured. The sweet-scented V. odorata 
is visited by the honey-bee, which insures cross-pollination, and in the absence 
of insect visits it effects self-fertilisation by cleistogamic flowers. The con- 
spicuous but scentless V. tricolor, var. i-ii'garis, is less frequently visited by 
insects (humble-bees). In V. silvatica and V. caiiina the pollen is carried on 

423. 421. -125. 

FIGS. 423-425. T~i'o(<i tricolor. 

FIG. 423. Capsule after dehiscence (nat. size). FIG. 421. External view of the seed. 

FIG. 425. Seed in longitudinal section. 

the head or proboscis of the honey-sucking bee. The fruits of F. odorata bury 
themselves slightly in the soil. In the others the fruits are raised above the 
ground ; the 3 boat- shaped valves close together along the central line, and 
eject the seeds, one by one, with much violence, so that they are thrown to a 
great distance. 

The Alsodeia-group has regular or almost regular flowers. Gamopetalous 
corollas are found in Paypuyrolea. Kativagesietf differs the most by its regular 
corolla, and 5- oo free or united stamiuodes. 

250 species ; especially in the Tropics. The lonidium-species are used as 
ipecacuanha. A number of Viola-species are cultivated as garden plants, 
especially V. odorata (sweet-scented Violet) and V. tricolor, which have a large 
number of varieties. 

Order 6. Frankeniaceae. A small order with doubtful relationships. 
Perennial herbs or shrubs ; beach plants with nodose stem. Sepals united, 
petals free. Unilocular ovary, with 3-4 parietal placenta?. Fruit a capsule. 
Embryo straight, eudospermous. Especially in S. Europe, Africa, on the 
shores of the Mediterranean and Atlantic. 

Order 7. Tamaricaceae (Tamarisks). To this order belong only Tamuri.v 
and Mtjricarla. They are shrubs of a cypress- or heather-like appearance, as 


the scattered leaves are very small, sessile, scale-like or linear, adpressed, 
entire, and usually glaucous, and the branches are slender and whip-like. The 
flowers are borne in small spikes or racemes, and are small, reddish or whitish, 
regular, $, hypogynous and polypetalous ; formula So, P5, Ao + (Tanuirix, 
which often has stipular teeth at the base of the filaments), or A5 + 5 (Myricariu, 
in which the stamens are united at the base) ; the number 4 may appear instead 
of 5, but in either case there is usually a tricarpellate gynceceum, which is 
iinilocular and has either parietal placent;e (\lyriearia) or a small basal 
placenta (Tamarix) ; 1 trifid style, or 3 styles. Capsule dehiscing along the 
dorsal suture, and resembling the Willows in having a unilocular ovary with 
numerous woolly seeds ; but the seed- wool in this case is b >rne on the chalaza, 
and may be attached to a long stalk. Some Tamarix-speeies shed part of their 
branches in the winter. 40 species ; North Temperate, on the sea-shores or 
steppes, especially in Asia. Ornamental shrubs: Myricana tjcnnanica, and 
Tama I'i.v ga llica. 

Order 8. Cistaceae. Shrubs or herbs, natives especially of 
the Mediterranean region. Flowers generally in raceme-like 
scorpioid cymes, regular, ^ > hypogynous ; sepals 5, free, twisted 
in the bud, of which the two outer are generally much smaller 
than the others; petals 5, free, twisted in the bud (in the direction 
opposite to the sepals), fugacious; stamens numerous; gynceceum 
syncarpous, carpels usually 8-5, style simple, ovary unilocular, with 
parietal placentation (seldom divided into loculi, with axile pla- 
centation). The ovules are orthotropous in opposition to some of 
the other orders of this family. The capsule dehisces along the 
dorsal sutures ; embyro curved. The leaves are simple, undivided, 
generally opposite and stipulate. They are Violaceaj with regular flowers, 
numerous stamens, and curved embryo. The numerous stamens are in reality 
only one or two 5-merous whorls, divided into a large number of stamens ; these 
are formed, therefore, in descending order, like the lobes of many compound 
foliage- 1 eaves. 

Helianthemnm (Rock-Rose), has 3 carpels. Cistus has 5 (10) 

About 70 species ; temperate climates, especially about the Mediterranean. 
The resin of the Cistas-species has been used medicinally (ladauum). 

Order 9. Bixaceae. This order is closely allied to the Cistaceas and 
Ternstroemiaceas ; like these it has regular, 5-merous, hypogynous flowers with 
numerous stamens, uuilocular ovary and parietal placenta; ; sometimes uni- 
sexual flowers ; it differs in having anatrcpous ovules, in the aestivation of 
the sepals, etc. All species (about 180) are trees or shrubs, with scattered, 
simple leaves, which usually have stipules, and are occasionally dotted with 
pellucid oil-glands. BLra ordlann (Trop. Am.) is the best known species; it 
has a 2-valved capsule; the seeds are enclosed in a shiny red, jlcslnj 
which contains the well-known orange or yellow dye, annatto. 



Order 10. Dilleniaceae. Gynceceum usually apocarpous, seed arillate. The 
flower has most frequently S5, P5, and compound stamens (one or more 
bundles); sometimes irregular. 200 species; Tropical; woody plants, many 
lianes. Dillenia, Camlollcn, Pleurandm, Dauilld, etc. 

Order 11. Elatinaceae (Water-worts). About 25 species belong to 
this order ; especially in temperate climates. They are small, creeping, rooted, 
aquatic plants, with opposite or verticillate leaves and stipules. The flowers 
are solitary or situated in small dichasia in the leaf-axils, they are small, 
regular, , hypogynous, with free petals, the same number in all 5 whorls 
(Sn, Pn, An + n, Gn), 3-merous (e.g. Elatine he.randrn), 4-merous (e.g. E. Jn/dro- 
jiipcf), or 5-ruerous (Bergiu) ; the corolla-stamens are sometimes suppressed ; 
petals imbricate without being twisted ; the ovary is 3-4-5-locular, with 3-4-5 
free styles ; the capsule dehisces septicidally. The seeds are orthotropous or 
curved, often transversely ribbed, endosperm wanting. The order is most 
nearly allied to Hypericaceaa, whose primitive form it appears to represent. 

FIG. 420. Diagram of Jfi/jii?nViti quadran- 
i/iilum: S indicates the bud of the helicoid 
cyme in the axil of the bracteole ft. 

FIG. 427. 
three bundles of stamens. 

Flower with 

Order 12. Hypericacese (St. John's-worts). This order 
is recognised by its always opposite or verticillate, simple, and 
entire, penninerved leaves, without stipules, and usually dotted 
with pellucid glands ; by the always $ , regular, hypogynous flowers 
in a cymose inflorescence ; the generally 5-merous calyx and 
corolla, with sepals and petals free ; the stamens 3-5, numerously 
branched (Figs. 426, 427) ; and the gyiioaceum, 3-5-carpellate, 
styles usually free. The ovary is 3-5-locular, or unilocular with 
3-5 parietal placenta?. Fruit a capsule (dehiscing septicidallv) or 
berry. Endosperm absent. 

The inflorescence is a dicJiasium or helicoid cyme. The structure of the 
flowers is the same as that of the foregoing orders : S5 , P5 ; succeeding these 
in some cases are two 5-merous whorls of stamens in regular alternation, of 


which the inner is epipetalous ; but the outer whorl is only represented by 5 small 
scales (Fig. 427), or is altogether absent (Hypericum catycinum, ]{. hircinum), 
and the inner divided into numerous stamens, that is, these 5 stamens are so 
deeply divided that 5 epipetalous groups bearing anthers are found (as in the 
Cistace.T) ; in other cases the flower becomes S-nierous alter the petal*, stamens 
3 + 3 following in regular alternation (Figs. 426, 427), the outer whorl of 
stamens in these cases is also present as staminodes (Fig. 427), or may be al- 
together suppressed. Carpels 3-5. The petals are often tivisted in the bud, and 
are then oblique. 

Hypericum. Some species have a square stem ; in these cases 
the leaves are placed opposite the edges. Fruit a capsule. Vismia 
has a berry. The flowers of Hypericum have no honey, and supply only 
pollen ; self-pollination often takes place. 

About '240 species ; the tropical ones being often shrubs or trees ; the others 
generally perennial shrubs. Hypericum, St. John's-wort, contains a resinous, 
red matter, which can be extracted with alcohol. The American gamboge is 
the dried sap of species of Vismia. 

Order 13. Guttiferae, or Clusiaceae. Closely allied to the Hypericacese 
and Ternstroemiaceaj. Leaves opposite or verticillate. The flowers are often 
unisexual ; stamens united ; the gynceceum has most frequently a sessile, 
radiating or shield-like stigma. 

370 species; chiefly in the Tropics (Am.). They are principally woody plants 
and their bark contains a yellow gum resin, " gamboge," which is extracted 
from Gurciiiia morella (E. Ind.) and others. Mangosteen (Garcinia mango- 
utaini S.E. Asia), and Mamniea americana (W. Ind.), have very delicious fruits. 
To this order also belong Platonia in*iy)ii*, Pentadesma liutyracea, (the Butter- 
tree), Clusid, Calophyllum, Cataba, etc. 

Order 14. Ternstroemiaceae. Trees and shrubs with 
scattered, simple, and often more or less leathery, evergreen, pen- 
ninerved leaves, without stipules (Fig. 428). The two most 
important genera are : Camellia and the closely allied Thea (by 
some authorities these are united into one genus). The flowers 
are regular, hypogynous, and situated singly on very short stalks. 
A number of green floral-leaves are placed below the calyx and 
gradually pass over into the sepals, and the leaves (5-6) of the 
calyx again gradually pass over into the corolla (this being especi- 
ally marked in Camellia), of which the number of leaves varies (5, 6. 
7 and upwards) ; the calyx and the corolla are acyclic or encyclic ; 
the petals are slightly united at the base ; stamens numerous in 
many whorls, the external ones are arranged in bundles and 
united with the petals as in the Columnifera? ; gynoeceum syn- 
carpous ; styles often free nearly to the base ; ovary 3-5-locular, 
ovules numerous in each loculus. The fruit is a woody capsule. 



Other genera show more distinctly than these the same structure as in the 
preceding orders, namely : So, P5, A5 + 5, of which the calyx-stamens are often 
suppressed, and the petal-stamens divided into numerous stamens. Kiclmei/i'i-n 
(S. Am.) 

260 species ; especially in the Tropics (E. Asia, Am.) The leaves of Thea 
chinensis (or Camellia thcti), the Tea-tree (E. Asia), are cultivated for the 
well-known " tea," and contain thein : the best are the young, still hairy 
leaves, of greyish colour; there are many varieties. Ornamental plants, 
Camellia japonica and Actinidia. 

FIG. 428. Thea chinensis (reduced). 

Closely allied to this order are : Order 15. Rhizoboleae (with enormously 
large hypocotyl hence the name), and Order 16. Marcgraviaceae (partly 
epiphytes, with dimorphic leaves and cup- or helmet-like, coloured, honey- 
secreting floral-leaves, which eerve to attract insects). 

Order 17. Dipterocarpaceae. This order has taken its name from the large 
wings attached to the fruits in Dipterocarpus (the wings being largely developed 
sepals) ; trees and shrubs from Trop. Asia. 180 species. Camphor ready 
prepared its found in the stem of Dryolalanops camphora. Hopea ; Valeria. 



Family 12. Gruinales. 

The flowers are hypogynous, $ , polypetalous, usually regular 
(except Pelargonium, Tropwolacene, Balsaminacese) and throughout 
5-merons : So, Po, A5 + 5, or 5 + 0, Go (eyripetalaus) . The stamens 
soon fall off and are obdiplostemonous, often united at the base 
(vionadelphous) ; the corolla-stamens are in some completely sup- 
pressed (e.g. Balsaminace&, Fig. 438), in others reduced to teeth 
(Linum, Fig. 431; Erodium). The Tropfeolacew have 3 carpels 
and only 8 stamens (Fig. 437). Ring-like nectaries are not pre- 
sent, but at most only glandular bodies, borne outside the base of 
the stamens. Ovaries many-locular. The ovules as a rule are 
pendulous, with the micro pyle directed outwards (Fig. 431, B), 
and the radicle therefore also points outwards. Usually herbs. 
Related to the Columniferae. 

Order 1. Oxalidaceae. Most of the species are herbs with 
rhizomes ; the leaves are stalked, compound, with entire leaflets 

which are folded and bent backwards in 
the bud (and in the sleep position), ex- 
stipulate ; some species have sensitive 
leaves. The flowers (Fig. 429) are re- 
gular, and have So, Po, which are twisted 
to the left or right in aestivation, Ao + 5, 
all united at the base (monadelphous), 
gynceceum 5-carpellate, styles 5 free, stig- 
mas capitate, ovary 5-locular, ovules 
numerous. The fruit is a capsule opening 
with clefts on the dorsal sutures through 
which the seeds are ejected, while the 

fleshy, external layer of the testa springs off elastically. Embryo 
straight. Endosperm. 

Oxalia (Wood-Sorrel). Leaves digitate. Species also occur with phyllodia, i.e. 
leaf-like petioles placed vertically without lamina ; a few have pinnate leaves. 
The flowers are situated singly or in dichasia, and unipared scorpioid cymes. 
The pollination is effected by insects. Some species are trimorphic (long-, 
short-, medium-styled flowers) and some, e.g. O. acetostlla, have cleistoganiic 
flowers in addition to the ordinary ones. Glands are found on the outer side of 
the corolla-stamens or of all the stamens. 0. tetrapliylla and others have 
adventitious edible roots, resembling tap-roots. Averrhoa is a tropical tree, with 
berries and pinnate leaves. 

235 species (205 belong to O.ralis); chiefly in S. Africa and Trop. America. 
Oxalate of potash is contained in the leaves of Oxalis. 

PIG. 429. Diagram of Oxal's 



Order 2. Linaceae. Hei-bs with scattered or opposite, sessile, 
simple, small, entire leaves, without (rarely with small) stipules. 
The flowers (Fig. 430) are regular, 5- or 4-merous. Petals are 
free, twisted, quickly falling off. Stamens united at the base ; the 
petal-stamens are either reduced to teeth (Fig. 431 A, in) or entirely 
suppressed. Styles free. The (5-4) epipetalous loculi of the 
ovary are incompletely halved by false divisional walls, each half 


430. 432. 

FIG. 430-433. Li'num usitatissimum. 


FIG. 430. The Flax plant. 

FIG. 431. A Flower after removal of sepals and petals ; m petal-stamens reduced to 
teetb. B Longitudinal section of ovary. C Transverse section of capsule. 

FIG. 432. Capsule (nat. size). 

FIG. 433. Transverse and longitudinal section of seed : 61 the cotyledons ; fc the 
plumule; R the radicle ; /) the endosperm ; s/c the testa. 

contains one ovule (Fig. 431 (7). The fruit is a spherical 
capsule, dehiscing along the divisional wall (Fig. 432) ; the 10 (-8) 
seeds have a straight embryo and very slight endosperm (Fig. 

W. B. 

E E 



Linum (Flax) has 5-merous flowers. The main axis terminates in 
a flower ; and the succeeding branching is cymose, or unipared scorpioid 
branching by unilateral development, and the flowers in consequence of the 
vigorous sympodial development of the lateral axis (and also by the leaves 
being displaced and pushed aside) , assume a position apparently lateral (i.e. 
racemose) without bracts ; each branch of the sympodium generally has 2 
leaves. The testa is shining and smooth when dry, but its external cellular 
layer becomes mucilaginous in water. Radiola has a 4-merous flower. 
It is a small herb with opposite leaves, and regular, dichasial 

The anthers and stigmas in L. catharticum and iisitatinsinnun develop 
simultaneously, and cross-pollination as well as self-pollination takes place. 
L. grandiflorum, perenne, and others, are dimorphic (short- and long-styled). 
There are 5 nectaries outside the stamens. 

130 species ; Linum and Radiola are native genera. L. usitatissimum is exten- 
sively cultivated in Europe (especially in Russia and Belgium), N. America and 
elsewhere (its home no doubt being Asia), partly on account of the oil (linseed 
oil) which is extracted from the seeds, and partly on account of the bast of the 
stem, which has very thick-walled cells. The seeds and oil are OFFICINAL. The 
species cultivated in ancient times was L. anrjiistifolium. Several species are 
cultivated as ornamental plants. 

Order 3. Geraniaceae. The majority are herbs with dichasial 
branching, and scattered or opposite, stalked, palminerced (rarely 
penninerved) leaves with small stipules. The flowers are regular 
(except Pelargonium) and 5-merous, with 10 or 5 stamens, which 
are slightly united at the base. Nectaries alternate with the 
corolla-stamens. The ovary is most frequently 5-locular, deeply 
5-grooved, and bears 1 well developed style (" beak "), which towards 
the apex divides into 5 branches bearing stigmas ; ovules 1 in each 
loculus, pendulous or ascending. The 5 carpels become detached 
from one another when ripe, and bend or roll back (Fig. 434) or 
become spirally tivisted in the upper " beak-like " part (Figs. 
435, 436), whilst a central column (septal column) pei'sists ; each 
carpel, in consequence, remains either closed, and the fruit is a 
5-merous schizocarp whose nut-like lower portion, containing the 
seed, is forced into the ground, thus burying the seed by the 
movements of the spirally-twisted, hygroscopic '' beak " (Figs. 
435, 436) ; or it opens along the ventral suture, so that the seeds 
may fall out, and it is then a 5-valved capsule, with septicidal de- 
hiscence (Fig. 434) and the rolling up often takes place so suddenly 
and violently that the seeds are shot out to considerable distances. 
The embryo is usually green and curved, and the cotyledons are 
folded ; endosperm is wanting. 



Geranium (Crane's-bill) lias 5 + 5 stamens, and a septicidal capsule; 
the carpels most frequently remain suspended from the apex of the 
column (Fig. 434). The leaves are most frequently palminerved. 
The flowers are situated solitarily or 2 together (2-flowered scor- 
pioid cyme). Erodium (Stork's-bill) ; inflorescence a many- 
flowered unipared scorpioid cyme, stamens 5 + (petal-stamens are 
wanting), and fruit a schizocarp whose carpels become detached; 
their beaks are hairy on the internal surface and twist themselves 
spirally (Fig. 436). The umbellate inflorescences are composed of 
multiflowered scorpioid cymes. The leaves are often penninerved. 
The most primitive type is represented by Biebersteinia: S5, P5, A5 + 5, Go 
(o varies free, and styles united above) ; fruit 5 small nuts. The most advanced 
type is Pelargonium, which has zygomorphic flowers, the posterior sepal being 

FIG. 431. Geranium san- 
guineum. Fruit (f). 

FIG. 435. Pelargonium. 

FIG. 436. E rod in m cica- 
tarium, detached carpel. 

prolonged into a spur which becomes adnate to the peduncle ; the petals are 
unequal in size ; some of the petal-stamens are often wanting. (Erodium may 
be slightly zygomorphic). 

POLLINATION. The large-flowered Geranium-species are protandrous, e.g. 
G. pratense (one whorl of stamens opens first, and then the other, and suc- 
ceeding these the stigmas, after shedding the pollen the stamens bend out- 
wards) ; the small-flowered are also adapted; with various modifications, for 
self-pollination. 470 species; moderately hot climates, especially S. Africa. 
Several Pelargonium-species, with numerous varieties, are ornamental plants 
(from S. Africa). 

Order 4. Tropaeolacese. Herbaceous, juicy plants which have 
scattered, long-stalked, peltate leaves without stipules, and often 



climb bj their sensitive petioles. The flowers are situated 
singly in the axils of the foliage-leaves on long stalks, and are 
zygomorphic, the receptacle under the posterior sepal being pro- 
longed into a spur ; there are also differences between the posterior 
and anterior petals, the 2 posterior petals situated on the border 

of the spur being perigynous, and the edge 
of the anterior petals adjoining the claw 
fringed. After the 5 sepals (which are 
more or less coloured) and the 5 petals, 
follow 8 stamens (as the 2 median ones are 
suppressed, one from each whorl) and a 
gynceceum formed of 3 carpels ; in each of 
the 3 loculi of the 3-grooved ovary is 1 
ovule. The fruit is a schizocarp and divides 
into 3 1-seeded, drupe-like fruitlets, which 
do not (as in the Geraniaceas) leave any 
pronounced column between them. Endo- 
The cotyledons are thick and sometimes 

FIG. 437. Diagram of 

Tropwolum: sp, spur. 

sperm is wanting. 

slightly coalescent. Tubers often occur. 

Tropceolum. About 40 species ; all from America. 

POLLINATION. The spur is the receptacle for the nectar ; the flowers are 
protandrous ; the anthers open first, and one by one take up a position in front 
of the entrance to the spur, resuming their original position when the pollen is 
shed ; the stigma finally takes their place after the filaments have bent back- 
wards. These plants Lave an acrid taste (hence the name " Nasturtium," 
" Indian Cress "), on which account the flower-buds and young fruits of T. 
inujus are used as capers. Some species are ornamental plants. 

Order 5. Balsaminaceae. Herbaceous, chiefly annual plants 
with juicy, brittle stems, so transparent that the vascular bundles 
may be distinctly seen. The leaves are simple, usually scattered, 
penninerved and dentate ; stipules are wanting, but sometimes 
large glands are present in their place at the base of the petioles. 
The flowers are strongly zygomorphic ; of their five 5-merous 
whorls the petal -stamens are suppressed (S5, P5, A5 + 0, Go); 
the sepals are coloured, the 2 anterior ones (Fig. 438 j, 5) are 
rri'i/ sniiiH. or entirely suppressed, the posterior one is very large 
and elongated into a spur, and the 2 lateral ones pushed forward ; 
sometimes the weight of the spur turns the flower completely 
round, so that the posterior leaves assume an anterior posi- 
tion ; apparently only 3 petals, since the lateral and the pos- 
terior petals become united in pairs, and the anterior is larger 


and differently shaped ; the 5 stamens have very short and thick 
filaments united at the base, and their anthers finally adhere to- 
gether and remain in this condition, covering over the gynceceum ; 
the filaments ultimately rupture at the base, and the entire 
anthers are raised on the apex of the gynoeceum as it grows up. 
The gyno3ceum has a sessile stigma and a 5-locular ovary. The 
fruit is a capsule which, on maturity, opens suddenly when 
irritated, dividing into valves from the base upwards, and as the 
5 valves roll up elastically, the seeds are shot out on all sides to 
considerable distances ; a central column persists (Fig. 439). The 
embryo is straight, and without endosperm. 

Impatiens ; in Europe only I. noli-me-tangere. 225 species; especially from 
Asia. Several species have two kinds of flowers: small, cleistogamic, but fertile; 

FIG. 433. Diagram of Impatiens glanduligera. FIG. 439. Fruit of Impatiens. 

and large, coloured flowers, which in I. lalsamine (ornamental plant, E. Ind.) 
are protandrous and pollinated by hive- and humble-bees, as they suck the honey 
from the spur. 

Order 6. Limnanthaceae. The flowers are regular and differ from all the 
other orders in the family by having the carpels not in front of the petals, but 
in front of the sepals (which are valvate), and further, the loculi are nearly free 
individually, but with a common (jynobaxic style ; the ovules are ascending aud 
apotroputts (auatropous with ventral raphe). The fruit is a schizocarp, with 
nut-like cocci. Limnanthes (4 species; N. Am.) perhaps belongs to another 

Order 7. Humiriaceae. Trees and shrubs; about 20 species; Trop. Am. 

Family 13. Columniferse. 

The chief characteristics of the orders belonging to this family 
are the $ , regular, generally 5-merous, hypogynous flowers with 


5-merous caly.v, sepals united and valvate in the bud; petals 5, 
free (often twisted in the bud); stamens oo e.g. : 10, in two whorls, 
but one of these is more or less suppressed, often altogether want- 
ing, or replaced by 5 staminodes, while tlie oilier (inner whorl) is 
generally divided more or less deeply into a large number of anther- 
bearing filaments. The filaments too (except Tiliaceee) are united 
into a tube, which, especially in the Malvacefe, forms a long column 
in the centre of the flower, surrounding the gyno?ceum (Figs. 445, 
448) ; in this case, which is the most pronounced, the filaments are 
united into one bundle (monad elphous), in other instances, poly- 
adelphous. The number of carpels varies greatly (2 to about 50), but 
they are neai-ly always united and form a syncarpous multilocular 
gynceceum. The vegetative characters also closely agree, the leaves 
are always scattered and generally stipulate; all the green portions 
very often bear stellate hairs, and the bark in all the 3 orders is rich 
in tough bast. Mucilage is often present in cells or passages. 
This family is connected with the Ternstroemiacese, from which it 
is very hard to draw a sharp line of demarcation, and it is also 
allied to the Cistaccse and to the Gruiiiales. 

Order 1. Sterculiaceae (including BiittneriacefE). This is, no 
doubt, the least modified order, and one in which the stamens occur 
undivided. Obdiplostemonous. The 10 stamens in two whorls 
are most frequently united at the base into a short tube, and have 
4-locular, extrorse anthers. The calyx-stamens are nearly always 
simple, tooth-like staminodes, situated on the edge of the tube, or 
are entirely suppressed. The same relation is found, for instance, 
in the Ampelidacea? and Rhamnacese, namely 5 stamens in front of 
the 5 petals; not infrequently the 5 stamens are doubled (Fig. 
441). Unisexual flowers are found in Sterculia, Cola, Heritiera. 
The corolla is often wanting, or developed in an unusual manner. 
Each loculus of the ovary (generally 5) always contains more 
than one ovule. Fruit a capsule. Androgynophore often present 
(Helicteres ; Sterculia, etc.). 

Hermannia, Mahernia, Melochia, etc., have flat petals with twisted aestivation ; 
5 undivided stamens, which usually are but slightly united at the base, and 
most frequentl.. without staminodes. Thoniania ; Hclicteres ; Sterculia (free 
follicles). Theou/otiij, Rulingia, Siittneria, Commersonia, Gunziuiia, etc., have 
petals concave at the base, and terminating in a limb abruptly bent back, and 
at the boundary between them most frequently ligular outgrowths, as in 
certain genera of the Caryophyllaceas ; stamens 5-15- <x, anthers at the edge of 
a short tube and 5 linear stamiuodes (Fig. 441). The Cocoa-tree (Theobroma, 
Fig. 440 bears large, reddish-yellow, berry-like fruits, resembling short cucum- 



bers, but ultimately becoming leathery to woody ; in each of the 5 loculi are 
2 (apparently only 1) rows of horizontal, oily seeds, as large as almonds, 
Cotyledons large, thick, and irregularly folded. Endosperm absent (Fig. 442). 
49 genera, with about 750 species ; almost entirely confined to tie Tropics ; 
none in Europe or in N. Asia. The seeds of the Cocoa-tree (T. cacao, bicolor, 
glaucion, etc., natives of Trop. Am., especially north of the Equator) are 

FIG. 410. Theobroma cacao. Branch with flowers and fruits 


411. A 412. B 

FIGS. 441-412. Theobroma cacao. 

FIG. 441. Diagram of the flower: st barren stamens. 

Fio. 442. B Seed in transverse section : ?i hilum. A Embryo after the removal of one 
of the cotyledons. 

used for chocolate and are also officinal (" Cocoa-beans," " Cocoa-butter," 
" Oil of Theobroma"). Theobromine. Cola acuminata, Africa. 

Order 2. Tiliaceae. This differs from the other orders of the 
Col umnif eras chiefly in the stamens being entirely free from each 
other, and also divided into many filaments, as far as the base, or at 
all events very far down, so that the flower appears to have numer- 



ous stamens or to be slightly polyadelphous (Fig. 443) ; in addition 
to this, it may be observed that the anthers are 4-locular and 
introrse. In Lilhea, the groups of stamens alternate with the 
petals. In a few genera (Corchorus, Triumfetta) 10 free and 
single stamens are found in 2 whorls ; but, in the majority, 
groups of free stamens in separate bundles. The stamens are 
more or less united in Apeiba, Lilhea. Style simple. Ovary 2- 

locular. The ovules are pen- 
dulous ; raphe turned inwards. 
The calyx readily falls off ; the 
festivation of the entirely free 
petals is slightly imbricate (not 

Tilla (Figs. 443, 444). Calyx 
and corolla 5-merous ; the 5 
staminal leaves (opposite the 
petals) divided as far as the base 
into a large number of stamens 
which are free or united into 
groups ; gynoeceum with 5 
loculi in the ovary (opposite the 
sepals) ; there are 2 ovules in 
each loculus, though the ovary 
ripens into a 1-seeded nut, which 
is not detached from the axis of 
the inflorescence, but is carried 
away by the wind, whirling 
round and round, its large- 
winged bracteole serving as a 
parachute (Fig. 443). Only trees, 
with alternate, obliquely heart-shaped 
and dentate leaves ; stellate hairs, as 
in the other Columuifera?, are often 
present. The terminal bud of the 
branch always fails to develop, and 
the growth is then continued sym- 
podially by the uppermost axillary 
buds. The INFLORESCENCE (Figs. 443, 444) is a 3-7-flowered dischasium (Fig. 
444 t, d, e), which is developed in the axil of a foliage-leaf (Fig. 444). The 
first of its 2 bracteoles (a) is large, thin, leaf-like, and united with the inflor- 
escence, the lower portion of which forms a broad wing, its so-called " bract " ; 
the second bracteole (b), on the other hand, remains scale-like, and supports 
a winter foliage-bud covered with bud-scales which thus is situated at the base 

FIG. 443. Inflorescence of Tilin, with its 
\vinged bracteole (h) ; a, a axis of the shoot; 
the vegetative bud is seen between the in- 
florescence and the axis of the shoot ; 6 
petiole of foliage-leaf. 



of the inflorescence, and is a bud of the 2nd order, in relation to the vegetative 
shoot. This bud is always found beneath the inflorescence on the branch 
placed horizontally, and the winged bracteole is always found above it, a relation 
which is connected with the fact that the 2 rows of shoots on the sides of a 
branch are antidromons with regard to each other. The dichasium itself (Fig. 
444) terminates with the flower (t) ; it has 3 floral-leaves (e, d, e), which soon 
fall off; c is barren: the other two bear flowers, or few flowered dichasia, or 
unipared scorpioid cymes (indicated in the figure). The folia?e-leaves are 
folded in the bud upon the median line (1, 2, 3 in Fig. 444 are foliage-leaves 
with their 2 stipules), the inner half is broader than the outer, and after unfold- 
ing is turned away from the mother-axis (the position of the new inflorescences 
and vegetative buds is indicated in their axils on the figure). The cotyledons 
on germination appear above the ground as large, I ibed leaves. 

Of the other genera some have a bell-shaped, gamosepalous calyx, some have 
no corolla, the anthers of some open at the apex (Aristotelia, Elaocarpus, etc.), 
the majority have a capsule, some have berries, or drupes, some separate into 
fruitlets, etc. Corchorus, Tiiumfetta (nut, with hooked bristles), Liihea, 
Apeiba,etc. Sparmannia is an African 
genus ; 4-merous flowers ; fruit a 
warted capsule ; filaments numerous 
and sensitive to touch, the external 
ones are without anthers and monili- 
forrn above. The plant is covered 
with numerous soft and stellate hairs, 
and at the apex of the branches bears 
several cymose umbels. 

POLLINATION in Tiliu is effected by 
insects, especially bees and Diptera, 
which swarm round the tree tops, 
allured by the numerous strongly- 

FIG. 444. Diagram of the inflorescence of 
Til in. and the vegetative bud ; the position 
of the leaves is indicated, and also the 
position of the inflorescences, which de- 
velop from their axils in the following 

scented flowers and the easily ac- 
cessible honey (formed in the hollow 
sepals). As the flowers are pendu- 
lous, the nectar is protected from 
ruin ; and, in addition, the inflor- 
escence is more or less concealed 
beneath the foliage-leaf. Self-pollination is impossible, on account of pro- 
tandry. About 470 species (nearly all trees and shrubs) ; especially in the 
Tropics, only a few being found in the temperate, none in the polar regions, or 
in high mountainous districts. The inflorescence of the native species of Tilia 
is medicinal. The wood is used for charcoal. The majority are used for timber, 
and for the sake of the bast (" Bast," " Jute," the bast of Co/chorus textilia, 
Liihea, and others). 

Order 3. Malvaceae (Mallows). The plants are easily re- 
cognised by the scattered, simple, palminerved, most frequently 
lobed, stipulate leaves, folded in the bud ; the perfect, regular, 
hypogynous flowers, with gamosepalous, persistent, 5-merous calyx 



with v air ate aestivation ; the 5 petals ticisfed in the bud and united 
with one another at the base, and by the 5 apparently numerous 
stamens (Figs. 445, 448), with the filaments united i>ito a tube, with 
renifurm btlucular anthers opening by a crescentic slit (in 2 valves). 
Carpels 3 oo united into onegynceceum ; the embryo is curredand 
the cotyledons are folded (Figs. 447, 451) ; endosperm scanty, often 
mucilaginous. Most of the plants belonging to this order are 
herbs, often closely studded with stellate hairs. The leaves are 
most frequently palmatifid or palmatisect. 

An epicalyx is often found formed by floral-leares placed close beneath the 
calyx, in some 3, in others several. The median sepal is posterior in the 
species without tpicalyx, often anterior in those which have an epicalyx. The 
petals are twisted either to the right or to the left in accordance with the spiral of 
the calyx ; they are most frequently oblique, as in the other plants with twisted 
corollas, so that the portion covered in the aestivation is the most developed. 
The corolla drops off as a whole, united with the staminal tube. Only the 5 
petal-stamens are developed, but they are divided into a number of stamens, 
placed in 2 rows, and provided only with /<Z/-anthers (leaf-segments, see Fig. 
446; the sepal-stamens are completely suppressed); these 5 stamiual leaves 
are then united into a tube, frequently 5-dentate at the top, and bearing the 

FIG. 415. Longitudinal section through the 
flower of Malva silvestris. 

FIG. 416. Diagram of Alilicei rosea : 
i the epicalyx. 

anthers on its external side. The pollen-grains are specially large, spherical 
and spiny. There are from 3 to about 50 carpels united into one gynceceurn and 
placed round the summit of the axis which most frequently projects between 
them. There is only 1 style, which is generally divided into as many stigma- 
bearing branches as there are carpels (Figs. 445, 448). The fruit is a schizo- 
carp or capsule. Endosperm (Figs. 447 A, 451) scanty, often mucilaginous 
round the embryo, which is rich in oil. 

The order is the most advanced type of Columniferffi ; it stands especially 
near to the terculiacere, but is separated from these and from the Tiliaceae, 
among other characters, by its 2-locular (ultimately 1-chambered; anthers. 

The sub-orders may be arranged as follows : 



I. Carpels in one whorl. 

A. The fruit a capsule, most frequently with loculicidal 
dehiscence, and many seeds in each loculus. 

I. GOSSYPIE^. The staminal-column is naked at the apex, 
blunted, or5-dentate. Gossypinm (the Cotton plant) has an epicalyx 
of 3 large ovate-cordate leaves, an almost entire, low and compressed 
calyx. Solitary flowers. Large, most frequently yellow, corollas. 
A 3-5-valved capsule with many spherical seeds. " Cotton " is the 
seed-hairs developed upon the entire surface of the seeds (Fig. 
447), and consists of long, 1-cellular hairs, filled with air (and 
therefore white) ; these are thin-walled, with a large lumen, and 
during drying twist spirally, and come together more or less in 
the form of bands. They consist of cellulose, and have a cuticle. 
Hibiscus has several, most frequently narrow, epicalyx-leaves, a 
distinct 5-toothed or 5-partite calyx. A^ntilon ; Modinla. 

A B 

FIG. 447. A Seed of Gossypium with hairs ; B the same in longitudinal section. 

2. BOMBACE^E. The staminal tube is more or less deeply cleft into bundles, 
sometimes almost to the base ; pollen smooth, style simple with capitate, lobed 
stigma. Almost all plants belonging to this group are trees, and in many 
instances have large barrel-shaped stems, that is, swollen in the centre, and 
sometimes covered with large warts. The wood is exceptionally light and soft. 
The flowers are often enormously large, and have beautiful petals ; iu some they 
unfold before the leaves. The capsule-wall is sometimes closely covered on its 
inner service with long, silky, woolly hairs, while the seeds themselves are gener- 
ally without hairs. These hairs, however, on account of their brittle nature, 
cannot be used like those of the Cotton-plant. Digitate leaves are found in the 
Baobab-tree (Adansonia) from Africa, noted for its enormously thick, but short 
stem, and in the American Silk-cotton trees (Bombax, Eriodendron, Chorisia). 
Ochroma, Cheirostemon, Durio, and others also belong to this group. Durio is 
noted for its delicious fruits, which have a most unpleasant smell. 

[Bombax malabaricum is diplostemonous ; the five sepal-stamens repeatedly 



branch, and the filaments bear unilocular anthers ; the five petal-stamens bear 
bilocular anthers.] 

B. Schizocarps, with 1-seeded fruitlets, most frequently nut- 
like and reniform (Figs. 449, 451). 

3. MALVE.E, MALLOW GROUP. The carpels are arranged in one 
whorl (Fig. 449) ; the number of stylar-branches equals that of 
the carpels ; fruitlets 1-seeded, reniform, indehiscent, but detach- 
ing themselves from one another and from the persistent central 
column (Figs. 450, 451). Malva has an epicalyx of 3 free lenves. 
A flower with 2 suppressed bracteoles is situated in the axil of the foliage- 
leaves ; one of these supports a homodromous foliage-shoot which forms a 
repetition of the main axis, the other an antidromous flower which continues 
the branching as a unipared scorpioid cyme. Althcea, Rose Mallow, has 
an epicalyx of 6-9 leaves itnited at the base.Lavatera, Sida, Anoaa, 
Bastardia, etc., have no epicalyx. 


448. 450. 451. 

FIGS. 448-161. Malva silvestris. 

FIG. 448. The flower after removal of the perianth (f). FIG. 449. The fruit (J). 

FIG. 450. A fruitlet (f). FIG. 451. The same in longitudinal section. 

4. URENK.E, have always only 5 carpels arranged in 1 whorl, with 1 ovule 
in each loculus, and the fruit a schizocarp, generally with nut-like fruitlets 
provided with warts and hooks ; but in some they dehisce by 2 valves (capsule). 
They differ principally from the other groups in baring twice as many sti/htr- 
Itranclies as easels ; the stamiual tube is naked at the point, blunt or 5 -toothed. 

-The genera Urena, Pavouia, Malachra, Malvavisais (with berry-like fruits) 
belong to this group. 

II. Carpels arranged in a spherical head in five groups opposite 
to the petals. 

5. MALOPF.E, differ from all the others in having a large number of fruitlets 
arranged irregularly in a round head, and separating considerably from each 



other even before maturity ; there is, however, only 1 style, divided into a 
corresponding number of branches (this condition may be considered to have 
arisen from the branching [dedoublement] of 5 carpels). Malope has 3 large, 
heart-shaped (Kitaibelia 6-9) epicalyx-leaves, united at the base. Palava has no 

POLLINATION. The majority have protandro is flowers, and are pollinated 
by insects. Between the basal portions of the 5 petals, there are 5 nectaries, 
protected from the rain by hairs, e.g. in Malva sUvestris. When the flower first 
opens the numerous anthers occupy the centre of the flower, and the still 
undeveloped stigmas are concealed in the staminal tube ; in the next stage the 
anthers are withered and empty, and the stigmas protrude and assume their 
places (Fig. 452). The large-flowered forms, it appears, are pollinated only by 
insects ; but self-pollination takes place in small-flowered forms, as, for example, 
in Malva rotundifulia, in which the stylar-brauches, twis'ing themselves, place 
the stigmas in between the undeveloped anthers. 

FIG. 452. Anodn liastata : a the bud just opened, the stigmas are concealed by the 
anthers; !> ffllly opened flower in ^ -stage; the upper stamens are developed first, and 
then the others in descending order; the stylar-branches are now visible, 'and lie bent back 
on the staminal column; c all the stamens project upwards, and all the anthers are open, 
bat the stylar-branches are still bent back; d the anthers are emptied and the filaments 
shrunk together, but the styles have now straightened themselves upwards, and the 
stigmas are in the receptive condition. 

DISTKIBUTION. 800 species (63 genera), most of which are natives of the 
Tropics, especially America. Althcea and some of the species of Malva are 
natives of the temperate regions of the Old World, the latter is also found in 
North America. Gossyjn'itwi is tropical, no doubt especially Asiatic (G. herbaceum 
from India ; G. arborenm from Upper Egypt). Cotton was introduced into 
Greece in the time of Herodotus, and was cultivated in America before the 
arrival of the Europeans. 

USES. Pungent and poisonous properties are entirely wanting ; mucilage, 
on the other hand, is found in abundance in all parts of the plant. Medicinal : 
the root of AWuca officinalis, leaves and flowers of Malva-species (M. silvestris 


7 nlgaris and lorealis) and Gossypiian. The seeds contain a large quantity of fatty 
oil, which is in some cases extracted (Cotton-seeds and others). The seed-liairs 
of the Cotton plant are the most important product of the order. The cultivated 
forms of Cotton belong to several species : G. barbadense, herbaceum, religiosum, 
arborenin (Nankin), hirsutum, and others. According to other botanists, there 
are only 3 species. Bast is obtained from e.g. Hibiscus cannabiinis (Gambo-hemp, 
Africa), Paritium tiliaceiim and Sida retuna. The fruits of certain species of 
Hibiscus (e.g. H. esculentu<, from Tropical Africa) are used in tropical countries 
as a vegetable before they are ripe. The colouring matter iu the flowers of 
Althtea rosea, var. niym, is used for colouring wines, and heuce is extensively 
cultivated in certain parts of Europe. Ethereal oils and sweet-scented flowers 
are rare ; but several species possess a peculiar musk-like odour (Malra moschata, 
Hibiscus abeluioschns, and others). Many are cultivated as ornamental plants 
on account of the large flowers, Hollyhock (A. rosea, etc.), Lacatera 
trimestris, Malope grandiflora and trijida, MaZca-species, Hibiscus rosa sinensi.*, 
syriaca ; Splueralcea, etc. 

Family 14. Tricoccae. 

The very large order Enphorbiacece and three smaller ones 
belong to this family. They have in common : unisexual, hypo- 
gynous, frequently regular flowers, the perianth most frequently 
single, rarely double, or entirely wanting ; there is such a great 
variety in the structure and parts of the flower that one only can 
be cited as the rule : viz. the simple gynceceum composed of 3 
carpels forming a 3-locular ovary, which is frequently more or less 
deeply grooved (hence the name, Tricoccce) ; in the inner angles 
of the loculi are found 1 or 2 (never several) pendulous (except 
-EWj9eraceoe),anatropous ovules, with upward and outwardly turned, 
frequently swollen, micropyle (Fig. 455). The seed most frequently 
has a large endosperm and a straight embryo (Figs. 455 B, 464). 
The family approaches the nearest to the Gruinales and Columniferre ; it may 
perhaps be regarded as an offshoot from the Sterculiaceas. 

Order 1. Kuphorbiaceae. Flowers unisexual. In each of 
the loculi of the ovary, generally 3, there are 1 or 2 pendulous 
ovules with upward and outwardly turned micropyle. The placenta 
protudes above the ovules (Figs. 4o4, 461 _#). On the ripening 
of the capsule the 3 carpels separate septicidally, frequently with 
great violence, ejecting the seeds and leaving a central column. 
Endosperm copious. For the rest, the flowers present all stages, 
from genera with calyx and corolla, to those which are the most 
reduced in Nature, namely the naked, 1-stamened flowers of 

The same variety which is found in the flower is also present in 


the vegetative parts. Some are herbs, as our Spurges, others are 
shrubs and trees ; some African Euphorbia-species even resemble 
the habit of a Cactus. Leaf-like branches with rudimentary leaves 
are found in Phyllantlms (sub-genus Xijloplnjlla) (Fig. 456). The 
leaves are scattered or opposite, often stipulate ; they are nearly 
always simple. Large, highly-branched cells containing a great 
quantity of pungent latex are found in many, and watery juice in 
others. Glands and glandular hairs are general. Only a few 
genera can be considered in this book. 

As an example of the most perfect flowers (which partly 
reproduce the Geraniaceous type) may be mentioned, Croton, 
Manihot, and Jatropha ; 5 sepals, 5 petals, sometimes gamopeta- 
lous, andrcecium diplqstemonous, or many-stamened, often mono- 



FIGS. 453-155. Ricinus commurus. 

FIG. 453. (J -flower (magnified). FIG. 454. -flower in longitudinal section. 

FiG.-455. A seed entire ; B in longitudinal section. 

Picinns (Castor-oil) (Figs. 453-455) ; monoecious; the ^-flowers, 
situated in the lower portion of the inflorescence, have 5 perianth- 
leaves and a large number of branched stamens ; the ? -flower 
has 3-5 perianth-leaves ; 3 bifid styles. Leaves peltate, palmately 
lobed. The seeds (Fig. 455) contain an abundance of fatty oil 
and large aleurone grains. Mercurialis (Mercury) : the perianth is 
most frequently 3-merous ; in the ^-flowers 9-12 stamens; in the 
$ -flowers most frequently a 2-locular gynoeceum. Phyllanthus : 
Pr3 + 3, A3, united in some and forming a column in the centre of 
the flower (Figs. 457, 458) ; Xylophylla is a section of this genus. 




Hura crepitans (Sand-box tree) has a many-carpellate gynceceum, 
which separates with great violence when ripe. A drupe is found 
in Hippomane mancinella (the Mancinil-tree, W. Ind.). Alchornea 
(Coelebngyne) ilicifolia is well known on accoiint of its "partheno- 
genesis "; only the ? -plant has been introduced into Europe, but it 
nevertheless produces seeds capable of germination; these have 
generally several embryos. 

Euphorbia (Spurge) has the most reduced flowers, which are 
-A. borne in a very complicated 

^v \ 

^$M& inflorescence. Each ^-flower 

(Fig. 460 B) is naked, and con- 
sists of one stamen only (ter- 
minal on the axis). In the 
closely allied genus Anthostema, 
a small perianth is situated at 
the place where, in Euphorbia, 
there is a joint in the "fila- 
ment," Fig. 461 A). The ?- 
flowers (Fig. 460) are naked, 
with a 3-locular ovary and 3 
bifid styles. (Anthostema has a 
distinct perianth (Fig. 461 B) ; 
in a few Euphorbias traces of a 
perianth are present). In Eu- 
phorbia the c- and ? -flowers 
are grouped into flower-like in- 
florescences termed " cyathia." 
Each cyathium consists of a 
centrally placed ? -flower which 
is first developed, surrounded 
by 5 groups of ^-flowers (sta- 
mens) placed in a zig-zag, with 
a centrifugal order of develop- 
ment (Figs. 459, 460 B\ that 
is, in unipared scorpioid cymes; 
these flowers are surrounded by 
an involucre of 5 leaves united 
into a bell-shaped stricture (Fig. 459, 1-5) (resembling a calyx) ; 
on its edge are placed 4, generally crescent-like, yellow glands, 
one in each of the intervals, except one, between the lobes of the 
involucre (shaded in Fig. 459; see also Fig. 460 A}. Scale-like 

-Plnjllanthus (Xylopltylla] 


FIGS. 45C-458. 

FIG. 456. Leaf-like branch with flowers 
(nat. size). FIG. -157. c? -flower ; and 
FIG. 458, ? -flower (mag). 


thin structures (floral-leaves ?)are situated between the ( -flowers. 
The ? -flower has a long stalk, and finally bends down on one side, 
namely to the place on the edge of the involucre where the gland 
is not developed. These cyathia are again arranged in an inflor- 
escence which commences as a 3-5-rayed umbellate cyme (pleio- 
chasium), the branches of which ramify dichasially and finally as 

FIG. -160. Euphorbia lathyris: A an (entire) inflorescence (cyathiutn) ; B the same after 
the removal of the involucre. 

FIG. 459. Diagram of an inflorescence (cyathmm) of FIG. 461. Anthostema . <J- 

Euphorbia with 3 floral-leaves, m,n, o, supporting other (4) and ?-(B) flowers; p the 

cyathia which are subtended by 2 floral-leaves (brae- perianth; ar the node; o the 

teolfs ; m, n). 1-5, the involucral leaves in their order ovule, 
of development; the shaded portions are the crescentic 

scorpioid cymes. Latex, with peculiar-shaped starch -grains, is 
found in laticiferous cells (especially in the Cactus-like, leafless 

205 genera ; more than 3,000 species ; especially in the Tropics. Many are 
used on account of the oil, and of the pungeut (aperient, poisonous, anthelmmtic, 
W. B. F F 



etc.) properties in the latex or the seeds. OFFICINAL: " Cascarilla-bark " of 
Croton eliitena ; the fatty oil of the seeds of Croton tiylium (Trop. Asia) ; " Cas- 
tor oil" from Eiciinis communis (Africa, aud cultivated in all warm climates 
throughout the world); the glandular hairs of Mallotits philippinensis (" Kamala ") ; 
this also yields a red dye. Gum " Euphorbium " is the hardened (resinous) 
latex of the Cactus-like Euphorbia resinifera (Morocco). NUTRITIVE plants : 
Jltiitihot utiliimima and other species (Maniok, Am.). Their large, farinaceous 
mots form a very important article of food in the Tropics (Cassava-flocr, 
Tapioca or Brazilian arrow-root). The fresh latex of the root in some species 
is a powerful poison ; but the poisonous properties are diminished by roasting 
or cooking. Caoutchouc is obtained from Siphonia elastica (Trop. S. Am.). 
The vegetable tallow of the Chinese tallow-tree (Stillingia sebifera) is used in 
large qautities in soap factories. An indigo-like dye is obtained from Crozo- 
plwra tiitctoria, and is also found in Mercnrialis pereuuis. Shellac is obtained 
from Alcurites laccifera, ORNAMENTAL plants: AcalypJia, Croton, Dalediainpia. 
- Hippomane is poisonous. 

Order 2. Buxaceae. This order differs from the Euphorbiaceffi in having 
the micropyle turned inwards ; the $ -flower has a 4-partite perianth aud 4 
stamens ; the $ -flower a 6-partite perianth and 3 carpels. Capsule with 
loculicidal deliiscence, the inner layer being detached elastically from the outer. 
30 species. Shrubs without latex and with evergreen leaves. Ihixus semper- 
rirens (Box) is an ornamental shrub (poisonous) ; it has a very hard and valuable 
wood which is used for wood engraving and carving. 

462. 463. 464. 

FIGS. 462-404. CalUtriche siagnalis. 

FIG. 462. <J -flower with the 2 bracteoles an<1 the solitary stamen. FIG. 463. $ -flower. 
FIG. 464. Longitudinal section of the ripe fruit. 

Order 3. Callitrichaceae. Aquatic plants, growing at the bottom of shallow- 
water, with opposite, simple, undivided, entire, exstipulate leaves, which are 
generally crowded and form a rosette in the apex of the branches. The flowers 
are unisexual (monoecious) and borne singly in the leaf-axils ; they have no 
perianth, but are provided with two delicate bracteoles ; the S -flowers consist 
of only 1 terminal stamen (Fig. 462) ; the $ -flowers of a bicarpellate gynoeceum 
(Fig. 463) which is originally 2-locular, but later on becomes 4-locular, as in the 
case of the gynceceuui of the Labiatae, by the formation of a false partition- 
wall ; in each loculus there is 1 pendulous ovule with the micropyle turned 
outwards. Fruit a ^-partite schizocarp (Fig. 464). 25 species. Callitriche. 

Order 4 (?). Empetraceae. 4 species. Empetrum ; E. >iirirum (Crowberry) 

is a heather-like, moorland, evergreen undershrub with linear leaves, having a 
deep groove closed with hairs, on the under side. The erect <>mles show the 
greatest deviation from the Euphorbiacere. Dioecious (and ) ; S3, P3 ; in the 
J -flower, 3 stamens; in the ? -flower, a 6-9-locular ovary. Fruit a drnj>,'. 

Family 15. Terebinthinae. 

The diagram of the flower (Figs. 455-467) is the same as in the 
Gruinales, namely S, P, A2 and G in whorls of 5 (less frequently 
3, 4, 6, 8), and the same modifications also occur with the suppres- 
sion of the petal-stamens, etc. But a rinj or sometimes cup-like 
glandular structure (disc) \sio\iudbetween the andrceciuni and the 
gynceceum (Figs. 465, 466). The flowers similarly are regular, 
hypogynous, $ and polypetalous, though exceptions are found to all 
these characters : thus, for example, united sepals and petals fre- 
quently occur, and, in some orders, unisexual flowers by the sup- 
pression of one sex. In most cases the flowers are small, greenish- 
yellow, and arranged in paniculate inflorescences. The carpels (most 
frequently 5) are free in a few, but generally united into a niulti- 
locular gynceceum ; rarely more than 1 or 2 ovules in each loculus. 
The gynceceum in the Anacardiaceas is so reduced that it has only 

1 fertile loculus with 1 ovule. The ovules are epitropous, i.e. ana- 
tropous with outward-turned raphe (except the Anacardiacea?). 
The majority of the species are trees and shrubs with scattered, 
often compound (pinnate) leaves without stipules, and as in addition 
they frequently contain aromatic, especially turpentine-like substances, 
they assume a certain resemblance to the Walnut trees, and were 
formerly classed with them mainly on this account. In a series 
of genera the volatile, scented oils are found in special glands in 
the bark of the branches and in the leaves, in the latter case 
appearing as pellucid dots. This family includes several orders 
which are somewhat difficult to distinguish from each other. 

Order 1. Connaraceae. This order forms the connecting link between 
Terebinthinas arid Rosiflor;e (.tymra) as well as Leguminosffi, with which they 
are sometimes classed. The flowers have 5 5-merous whorls; 2 ovules in each 
loculus; micropyle turned upwards. Fruit a fUirle, rarely a collection of 
follicles. Seed with aril. Shrubs with scattered (most frequently pinnate) 
leaves, without stipules. 170 species. Tropical. 

Order 2. Meliaceae. Trees and shrubs with scattered, often pinnate leaves 
without pellucid dots and exstipulate ; the leaflets are nearly always entire. 
Flowers small in paniculate inflorescences. Calyx and corolla 4-5-merous ; 

2 whorls of stamens ; 3-5 carpels in the gynceceum. A very characteristic 
feature is the union of the filaments into a tube, on the edge of which stipnlr- 
like teeth are often found. There are most frequently 2 ovules in the loculi ; 



fruit a capsule with many winged seeds in Swietenia (Mahogany tree ; Trop. 
Am.), C'edrela, etc. ; berries in others. The wood of Cedrela is used for making 
cigar boxes. 550 species ; tropical. 

Order 3. Rutaceae. Leaves glandular with pellucid dots. 
The type is the same as that of the family. Flowers 4-5-merous. 
The ovary is most frequently 4-5-grooved. Disc well pronounced, 
often appearing as a "gynophore." The majority are shrubs with 
alternate or opposite, compound, more rarely simple, leaves. 

A. The ovary is deeply 2-5-cleft with basal styles which are 
more or less united ; the carpels in some genera are entirely free 
(groups 1, 2). The fruit is capsular and most frequently dehisces 
like follicles along the ventral suture or septicidally, so that a 
horn-like internal layer (endocarp) separates elastically from the 
external layer. 

1. ZANTHOXYLE.E. ZantUoxylum ; Choisya ; Evodia. 

2. BOKONIE.E. Australia. Correa. 

3. DIOSME^;. Heather-like shrubs; Africa. Diosma, Coleonema, En>j)leimnn 
and Barosma. OFFICINAL : Baro*ma crenulata and betulina, " broad Buchu 
leaves" (B. serratifolia and Empleurum semtlatum, "narrow Buchu -leaves"). 

FIG. -165. Enta. Flower (mag.). 

FIG. 466. Eula. Longitudinal section of flower. 

FJG. 467. Ruta. Floral diagram. 

4. RUTEJ:. Ruta (Figs. 465-467) graveolens is an herbaceous, 
glaucous, strongly smelling plant with bipinnate leaves and yellow 
flowers ; the terminal flower is 5-merous, the others 4-merous 
(S. Eur.). Dictamnus ; zygomorphic flower. The individual carpels 


of the fruit separate from each other, and dehisce like follicles, upon which the 
internal layer is detached elastically and springs out, carrying the seeds with it. 
Several species are ornamental plants. 

5. CUSPARIE.E. American. Flowers often zygoinorphic with garnopetalous 
corolla; stamens 5. Ticorea ; Galipea (G. oflicinalis ; S. Am. ; "Cortex an- 
gosturas ") ; Cusparia; Almeidea. 

B. The ovary is entire or only slightly grooved ; the style is 
terminal, undivided. The fruit is most frequently a drupe or 

6. TODDALIE.E. Ptelea ; winged fruit. The huds are enclosed in the leaf- 
sheath. Skimmia ; Phcllodendron. 



FIGS. 468-170. Citrus vulgaris. 

FIG. 468.-Branch with compound leaves. FIG. 4<59.-Transverse section of fruit. 

FIG. 470. Flowers (after the removal of the petals). 

7. AURANTIEJ:, ORANGE GROUP. Fruit a berry with a leathery 
external layer. The most typical flower is found for example in 
Limonia: S5, P5, A5 + 5, G5 (2-5). Citrus has 4-3-8-mevoiis 


flowers, a gamosepalous, dentate calyx, free petals, one whorl of 
stamens which are split irregularly into several bundles (Fig. 470). 
The fruit is a multilocular berry provided with a thick, tough, outer 
layer. The juicy pulp, which fills up the loculi and envelopes 
the seeds, is formed from many large-celled, juicy hair-structures 
which arise on the inner side of the walls of the loculi and by 
degrees entirely fill them up ; the dissepiments remain thin, and 
form the partitions so easily separating from each other (Fig. 
469). The seeds in many instances are remarkable for containing 
several embryos. The blade of the leaf is separated from the 
frequently winged stalk by a node (and hence is a compound leaf 
with only the terminal leaflet developed ?) (Fig. 468) ; in other 
genera, as Triphasia, there is a fully developed trifoliate leaf. 
Thorns are frequently developed. The species of this geuns, which is 
a native of the warmer parts of S. E. Asia, are very hard to separate. The 
differences are found in the foims of the fruit, the leaves and the leaf-stalks, 
and in the number of stamens. Citrus medica, " Cedrat " (Ind.) ; C. linionuiit, 
"Citron," "Lemon" (introduced into Italy in the 3rd to 4th century). 
OFFICINAL: the fruits and essential oil of Lemon. C. aurantium from E. Asia, 
the Orange (introduced into Italy in the 14th century). C. vulgaris (Fig. 468), 
Bitter Orange (introduced into Europe at the time of the Crusades) ; the unripe 
Bitter Oranges, and peel of the Bitter Orange is officinal ; it is from the flowers 
of this species especially that the essence of Neroli is made. (.'. limetta, 
C. bergamia, Berganiot ; essence of Bergamot is officinal. C. decumana, Pornalo, 
a native of the Islands of the Pacific. About 780 species ; chiefly tropical. 

Order 4. Burseraceas. Fruit a drupe ; 1-5 stones. The bark, as well as 
the other parts, contain strong aromatic resins and balsams, and hence 
several species are used : the Myrrh tree, Commiphora (Balsamodendron) from 
Arabia and Africa; OFFICINAL: Myrrha (Cominipliora mi/rrha). Mecca-balsam 
from C. opobalsamum, Arabia ; E. Afiica. The Incense-tree (Soswellia) from the 
same parts of the globe and E.India. The incense of B. carteri is medicinal 
(Frankincense). The resin (Elemi) of Protiitm-species is officinal, and is used 
technically for varnish (S. Am.j. Takamahaka- resin from Elapltrium (S. Am.) 
Prolium (Icica) ; Annjris (1 carpel). 270 species ; tropical. 

Order 5. Zygophyllaceae. The majority have opposite, pinnate leaves with 
stipules. Leaves without pellucid dots. The filaments have a scale on the 
inner side. The most important is Guaiacum qfficiunle (West India), the wood 
(Lignum Vitas) of which is very hard and heavy, this wood and Gum-guaiacum 
are officinal. Others have a peculiar repulsive smell and taste : the Creosote 
shrub (Lama mexicana) and Zygopltyllum simplex. Tribiilus ten-ester is a 
common weed in S. Europe. Fagohia. Peganum Imrnnila (South of Russia) 
yields a red dye. 110 species ; especially iu the Tropics ; several species iu 
sandy deserts. Nitraria. 

Order 6. Simarubaceae. This order is distinguished by the abundance of 
liitter substances which it contains (Quassiue) especially in the bark and the 
wood. The wood of Quassia amara (Guiana, Antilles) is officinal; Picraena 


I'.rci'lsd yields Jamaica Quassia; the bark of Simaruba, Simula-species and 
others is used. Ailanthm glandulosa is a garden plant (pinnate leaves, winged 
fruit). 110 species. Tropical. 

Order 7. Ochnaceae. Flowers diplosternonous, 5-merous. The unilocular 
ovaries, which are individually free, project considerably into the air around the 
gynobasic style ; 1 ovule in each loculus ; the fruitlets are drupes. Shrubs ; 
leaves alternate, with stipules. Ochna ; Ouratea. 160 species; tropical; 
especially American. 

Order 8. Anacardiaceae. The ovary rarely contains more than 1 ovule, 
even though there be several loculi and several carpels; in Anncardlnin all the 
10 stamens except one become suppressed. Eesin passages. Anacardium. The 
most peculiar feature is the development of the flower-stalk into a fleshy body 
about the form and size of a pear (A. occidentale from Trop. Am. and A. 
orientate from E. Ind.) which bears the kidney-shaped nut (the so-called 
"Cashew-nut") on its apex. Manylfera indiai (the Mango-tree, from E. 
Ind.) is cultivated in several tropical countries on account of its delicious 
drupe. Similarly, species of Spoiidias (S. dulcis, Pacific Islands, S. Ititea). 
Several species of Kims are ornamental shrubs in this country, for instance, 
R. typhina (N. Am.), R. cotiiius (the Wig-tree, the barren flower-stalks of the 
panicles being feather-like and hairy) ; R. toxicodendron (Poisonous Sumach, from 
N. Am.) is poisonous. Chinese galls are produced by the sting of a leaf-louse 
(Aphis chinensis) on R. semialata (China), and Japanese wax is from the the 
seeds of R.succedanea (Japan). Considerable quantities of Sumach (R.coriaria) 
are used in tanning and as a black dye. OFFICINAL : the mastic resin of 
Pittacia lentiscus (the Mastic-tree, from the Mediterranean). The fruits of 
Piitucia vera (Syria) are edible; P. terebinthus and others yield turpentine. 
450 species ; tropical. 

Order 9. Icacinaceae. Flowers 4-5-merous ; haplostemonous ; receptacle 
convex or cup-like surrounding the gynceceurn ; in the (single) loculus of the 
ovary, 2 anatropous, pendulous ovules. 200 species ; tropical. 

Family 16. Aesculinae. 

The essential characters of this family are in the main the 
same as those of the Terebinthinas and Gruinales. The flowers are 
hypogynous, perfect, with free petals, 5-merous (S5, P5, typically 
A5 + 5, all of which, however, are not generally developed ; in our 
native orders there are only 7-8 stamens), and most frequently 
a 3-meros, 3-locnlar gynceceiim (less frequently 2 or 5 carpels with 
as many loculi). In each loculus there are usually only 1-2 
ovules. A deviation from the preceding families is the frequent 
zygomorphy of the flower, with, as a rule an oblique plane of 
symmetry (Fig. 471). When a disc is developed it is placed 
outside the stamens. The majority have no endosperm (Fig. 473). 
The members of the family are nearly all trees. 

The family is closely allied to the Terebinthinfe, but unlike this it never has 



aromatic properties, and differs also in the position of the nectary, in the 
flowers, which are often irregular with a reduction in the number of stamens, 
and in the ovule which is usually ascending with micropyle pointing downwards 
(the Terebinthinae having the micropyle turned upwards), etc. It is also related 
to Frangulinaa, the Staphyleaceae being the chief connecting link ; but the 
.^Ssculinas generally have compound leaves. 

Order 1. Staphyleaceae. Leaves opposite, often compound. Flowers 
regular, $ > 5-merous in calyx and corolla, 5-stamened. The stamens are placed 
outside the nectary. Ovary syncarpous or 2-3-partite with free styles. The 
capsule is thin, bladder-like, 2-3-locular, opening at the apex, and has several 
very hard seeds with a shining testa without aril. Endosperm. Staphylea 
pinnata (S.Europe) and trifoliata (N. Am.) are cultivated in gardens; they 
have white flowers in pendulous, axillary racemes or panicles. 16 species. 
Staphylea is found in the Tertiary of N. America. 

Order 2. Melianthaceae. Glaucous shrubs with scattered, pinnate leaves, 
and large stipules. Melianthus. 8 species ; S. Africa. 

Order 3. Sapindacese. Trees or shrubs, often climbing by 
tendrils (lianes with anomalous structure of the stem) and with 
compound leaves. The flowers, in most cases, are small, insigni- 

471. 472. 473. 

FIGS. 471-473. JEsculus hippocastanuin. 

FIG. 471. Diagram of the flower and of a scorpioid cyme. FIG. 472. Flower in 
longitudinal section. FIG. 473. Seed in longitudinal section. 

ficant, and without scent, and in some polygamous and zygo- 
morphic. S4-5, P4-5, A8 (less frequently 5-10) inside the nectary 
(disc) ; ovary generally 3-locular, with 1-2 ovules in each loculus 
(raphe ventral, micr-opyle turned downwards). Seed without 
endosperm, often with an aril. The embryo is often thick and 
curved (Fig. 473). 

JEsculus (Horse-Chestnut). Trees with opposite, digitate, den- 
tate leaves without stipules ; the inflorescence is composed of 
unipared scorpioid cymes arranged in a pyramidal panicle (termed 
a thyrsus). The flowers are irregular, with an oblique plane of 



symmetry (through the 4fch sepal, Fig. 471); there are 5 sepals, 5 free 
petals, of which the one lying between S 3 and S 5 is the smallest 
(see Fig. 471) and may be absent ; stamens 7 (5 + 2), three being 
suppressed ; gyncecenm simple, 3-carpellary and 3-locular, with 
single style ; of the two ovules one is ascending, the other descend- 
ing (Fig. 472). The fruit is a 3-valvate, sometimes spiny, capsule, 
with loculicidal dehiscence, the seed having a large hilum, a 
curved embryo without endosperm and united cotyledons (the 
radicle lies in a fold of the testa, Fig. 473). JR. Tiippocastanum 
(Greece, Asia), introduced into cultivation about 300 years ago ; 
the majority of the other species, e.g. 2E. pavia, etc., several of 
which are frequently cultivated in gardens, are from N. America. 
The flower of the Horse-Chestnut is adapted for bees, whose abdomen touches 
the anthers or style when visiting the flower. The flowers are protogyuous. 

The other Sapindacefe have most frequently 4 sepals, 8 stamens, various 
fruits (septicidal capsule, nuts with or without wings, schizocarp), etc. Ser- 
jania, Cardiospermum, Sapindug, Koelreutiria, etc. (about 118 genera, 970 
species). The seeds of Paullinia sorbilis contain caffeine, and are used as 
"Pasta guarana," in the North Western Brazils in the manufacture of a 
common drink. Nephelium (or Euphoria) litchi (with edible aril), and other 
species, from Asia. 

Order 4. Aceraceae. This order is so closely allied to the 
Sapindacea?, that some authorities have classed it with them. 
The main difference is 
in the regularity of the 
flowers, and the 2-mer- 
ous gynoeceum (in ab- 
normal cases several 
carpels occur). They 
are trees, and, like the 
Horse- Chestnuts, have 
opposite leaves without 
stipules ; in Acer the 
leaves are palminerved, 
but imparipinnate in 
Negundo, a plant fre- 
quently cultivated in 
gardens. The flowers 
are often unisexual, polygamous (some species have $-,<?- and 
? -flowers); sepals 5, petals 5 free, stamens 8 (that is, 5 + 5, but 
the two median ones are absent) inside a large disc. Fruit a 
samara (schizocarp) with 2 winged, nut-like fruitlets (Fig. 474). 

FIG. 474. Samara of Acer platanoides. 



In each of the 2 loculi of the ovary are 2 ovules. Embryo curved, 
with thin, folded cotyledons. Endosperm absent The inflorescences 
are racemes with a more or less elongated rnaiu axis and terminal flower (which 
sometimes has 10 stamens); when the lateral branches are developed they are 
similar to the main axis. In some species both corolla and petal- stamens are 
suppressed. Acer is pollinated by insects, Nee/undo by the wind. 88 species ; 
North Temperate zone. Acer in the Tertiary from the Oligoceue. The follow- 
ing are native plants: Maple (Actr campestre), Sycamore (A. pseudoplatanus, 
doubtful native). Important as avenue trees and timber. Sugar is obtained 
from the spring sap of the Sugar Maple (N. Am.). 

Order 5. Malpighiaceae. A tropical (especially American) order closely 
related to the Aceraceae, having often the same form of fruit (but 3-partite). 
Some species are lianes with anomalous stem-structure Leaves opposite. The 
flowers are regular or obliquely zygomorphic (the plane of symmetry passing 
through sepal 3, with So, Po, A5 + 5, G3 ; 1 pendulous ovule in each loculus. 
Important characteristics for identification are the numerous graudular struc- 
tures on the sepals. Peculiar 2-spiued hairs are found in some. Malpigltia, 
Bunchosia, Galphlmia, Tetnipterh, Heteropt<ris, etc. About 600 species. 

Order 6. Erythroxylaceae. Sepals 5, petals 5 (with a ligular corona), 10 
stamens in one bundle. Gynceceum 3 locular. Fruit a drupe. Tropical 
(especially American) trees ami shrubs, the Coca-plant (Erythroxylon coca) 
being best known. Its leaves are considered by the inhabitants of Chili and 
Peru to be one of the indispensable necessaries of life ; they are chewed, and 
possess intoxicating, exhilarating properties, and contain the alkaloid cocaine, 
which is frequently employed as a local anaesthetic. 103 species ; chiefly in 

Vochysiaceae. Trees ; Trop. Am. 1 stamen. 140 species. 

Trigoniacese. Shrubs; Trop. Am. 30 species. 

Tremandraceae. Polygalaceaa with regular flowers. 27 species. 

Order 7. 

Order 8. 

Order 9. 


Order 10. Polygalacese. Herbs or shrubs (some tropical 
species are lianes) with scattered (rarely opposite), simple a,nd 
most frequently quite entire leaves, without stipules. The flowers 

are usually borne in terminal spikes or 
racemes, and are strongly zygomorphio 
(the plane of symmetry being median) ; 
they have 5 free sepals, the 2 lateral 
ones of which (4 and 5 in Figs. 475, 47b') 
are very large, petaloid, and frequently 
project on each side like the "wings" 
of a Pea- flower ; petals 5, of which the 
two lateral ones are wanting or rudi- 
FIG. 475. Diagram of Poly- mentary (dotted on Fig. 475), and the 

gala : d a gland in the posterior anter i or the keel " (Fig. 476 c) is large, 
side of the flower; a and the 

two caducous bracteoies. hollow and boat-shaped, and frequently 



with a lobed or fimbriated edge (Fig. 476 .1 and L, c) ; stamens 8, 
the two median ones being absent, all united into a tube split 
along the back, which is also slightly united to the keel (the 
anthers, often 2 locular, open by pores, Fig. 476 Z?, st) ; the '2 median 
carpels form a bilocular ovary. 1 pendulous ovule in each loculus 
(Figs. 476 C, 475) ; capsule compressed with loculicidal dehis- 
cence, rarely a nut. Poly gal a (Milk- wort). 

470 species ; distributed over the whole globe (none Arctic). OFFICINAL : the 
root of P. smga, from N. Am. Some are used as ornamental plants. 

POLLINATION. The flowers of Polygala are pollinated by insects (chiefly bees). 
The rimbriated processes of the anterior petal support the insect when it alights. 
The anthers lie on each side of the stigma in the pouch of the anterior petal ; 
the apex of the style is spoon-shaped, and immediately behind it is a viscid 
stigmatic lobe. In reaching the honey the proboscis of the insect must come 
in contact with the pollen and the viscid stigma, by which it is rendered sticky ; 
this ensures the pollen adhering to the proboscis and so being carried to other 

FIG. 476. Polygala amara. Parts of the flower (mag.) A Flower from side, 1-5 sepals : 
c keel; B flower from above spread out : st the 8 stamens ; o fimbriated edge of "keel"; 
(.' ovary with style and stigma. 

Family 17. Frangulinae. 

The plants belonging to this family, with very few exceptions, 
are trees or shrubs. The leaves are usually simple ; stipules may 
be absent or present. The flowers in almost all the orders are 
small, green or whitish; they are always regular, 4- or 5-merons with 
2-5 carpels, but never have more than 1 whorl of stamens, which 
in Ehamnacece and Ampelidacece are placed opposite the petals 
(typically 5 + 5 or 4 4-4 stamens, of which however either the ex- 
ternal or internal whorl is always wanting) ; hypogynous or 
slightly perigynous, in Rhamnacew only strongly perigynous or 
epigynous ; generally ^ ; the calyx is inconspicuous ; petals free or 


slightly united. Gjnoeceum simple ; ovary generally multilocular ; 
style short or entirely wanting. A disc is nearly always developed 
in the flower, but is found sometimes inside the staminal whorl, 
sometimes outside it or between the stamens. The ovules are 
apotropous (anatropous with dorsal or ventral raphe). 

Order 1. Celastraceae. Euonymus enropcea (Spindle-tree) may 
be chosen as a type. It is a shrub with simple, opposite leaves 
and small caducous stipules. The small, greenish-yellow flowers, 
borne in regularly-branched dichasia, are regular, ^ , with 4 whorls, 
4- (or 5-) merous in regular alternation. There is a thick disc 
upon which the polypetalous corolla (imbricate in the bud) and 
the stamens are borne, with a slightly perigynous insertion. The 
style is short and thick ; the ovary has 2 erect ovules in each 
loculus. The fruit is a red, 4-valvate capsule with loculicidal 
dehiscence ; the seeds are few in number, and have a large, red- 
yellow aril (developed from the micropyle). Embryo green, in a 
large, fleshy, white endosperm. The dingy yellow flowers are generally 
visited only by flies and ants for the sake of the honey secreted by the disc, and 
while they run about on the flowers they touch the anthers and stigmas, now 
with one part of the body, now with another. The flower is protandrons. The 
stigmas are not developed till several days after the opening of the anthers. 
Celastrus, Cassine, Catha, etc. 

38 genera ; 300 species. Distributed over the entire globe, with the exception 
of the colder districts, and especially in the Tropics. Some are ornamental 
bushes (Ruonijmm japoitica). The leaves of Catha edulis are used by the 
Arabs and Abyssinians in the same way as those of Coca by the Peruvians. 

Order 2. Hippocrateaceae. 150 species; tropical; chiefly lianes. So, 
P5, A3, G3. Anthers extrorse. 

Order 3. Aquifoliaceae (Hollies). The genus Ilex forms 
almost the entire order. (1/5 species out of 180 ; especially from 

S. Am.) They are shrubs or trees 
with scattered, leathery, simple 
leaves (in Ilex aquifolium, spiny) 
with very small stipules. The 
flowers are small, white, and borne 
in few-flowered inflorescences in the 
axils of the foliage-leaves ; they are 
most frequently unisexual and dioe- 
cious. There are 4-5 sepals, petals, 
stamens and carpels in regular al- 
ternation ; the calyx and corolla 

FIG. 47". Ilex aquifolium : .. . , 

magnified flower. have their leaves slightly connate ; 


stamens slightly adnate to the corolla; the ovary is generally 
almost spherical with a thick, sessile stigma (Fig. 477). This 
order deviates especially from Celastraceas in the absence of the 
disc and in having only 1 (pendulous) ovule in each of the 4 
loculi of the ovary, and in having a drupe, with generally 4 stones. 
Embryo extremely small, at the apex of the large endosperm, with 
the radicle directed upwards. 3 genera. I. aqm folium (Holly) principally 
on the coasts of European countries; from Norway toW. Denmark, and further 
westward. It is a common garden shrub with stiff, shining leaves and red fruits. 
Several South American species contain so much caffeine that they may be used 
as a beverage in the place of tea (I. paragitayensis, Paraguay tea, or Mate). 
The Holly does not contain caffeine. 

Order 4. Ampelidacese (Vines). Shrubs with the stem 
swollen at the insertion of the petioles and climbing by tendrils 
borne opposite the leaves (Figs. 478, 479). The leaves are scattered 
(generally |), stalked, stipulate, frequently palminerved and lobed, 
divided or compound. The small, greenish flowers are generally 
borne in paniculate inflorescences, whose position is the same as that 
of the tendrils (Fig. 478) ; they are hypogynous or slightly peri- 
gynons, $, with 4-5 sepals, petals, stamens (which, as in the 
Rhamnea3, are opposite the petals ; Fig. 480 A, B) and 2 carpels. The 
calyx is very small, entire, or slightly dentate ; corolla valvate, and 
in some falling off as a hood, since the individual parts remain 
united at the summit (Fig. 480 A). Between the stamens and 
gynceceum is situated an hypogynous disc, with 5 lobes alternating 
with the stamens (Fig. 480 A, _Z>, E). In each loculus of the 
2-locular ovary there are 2 erect ovules (E) ; the style is short or 
wanting. The fruit is a berry. The embryo is small and lies in 
a horny, sometimes slightly folded (ruminate) endosperm (Fig. 
480 C, D). 

Vitis and Ampelopnis (5-merous flowers) ; Cissns (4-merous flower) ; Leca 
(without stipules, corolla gamopetalous). The inflorescence in Pterisanthes 
(E. Iiid.) has a peculiar, flat, leaf-like axis, on the edges of which $ -flowers 
are borne, and on the surface $ -flowers. 

The TENDRILS in Ampelidacese are modified branches, since they bear leaves 
and may be abnormally developed as branches with foliage-leaves, and finally 
the inflorescences are borne in the position of the tendrils, and tendrils are met 
with which are partly inflorescences. The explanation of the position of the 
tendril, namely, right opposite the foliage-leaf but without a subtending-leaf, 
has been much disputed. The relative positions are as follows : in Vitis vi it if era 
the following two kinds of shoots and relative positions are found (the other 
species deviate in one or other particular), (a) LONG-BRANCHES, which have 
2 scale-leaves and a large number of foliage-leaves with a divergence of i ; 



opposite the lowest 3-5 foliage-leaves no tendrils are found, then follow : 2 
foliage-leaves with tendrils, 1 without a tendril, 2 with and 1 without, etc., 
with great regularity. Buds are developed in the axils of the foliage-leaves 



479. 481. 

Ftcs. 478-181. Vitis vinifera. 

FIG. 478. Branch with bunch of grapes. 

FIG. 479. Diagram of the position of leaf and tendrils. The branch is divided into 
sections on the sympodial theory (the successive generations, I, IT, III, IV, are alternately 
white and shaded) ; fc buds. 

FIG. 430. A Flower throwing off the corolla ; B flower after the removal of the corolla ; 
C, D longitudinal and transverse section of seed ; E longitudinal section of gynoeceum ; s 

FIG. 481. Diagram of branch ;md position of leaves; si tendril; It the main axis; ox 
stipules of the foliage-leaf shown below; y axillary -bud (the dwarf-branch); v its fore- 
leaf; !, Kits first two foliage-leaves with their stipules; lt l long-branch in the axil of v 
(everything appertaining to this branch is entirely black) ; r x the first leaf of this branch. 


(Fig. 479): these develop into (l>) DWARF-BRANCHES, which commence with 
1 laterally-placed scale-leaf (fore-leaf ; Fig. 481 v) succeeded by several 
foliage-leaves with a divergence of J (in a plane at right angles to th it of the 
mother-shoot), but the whole shoot is extremely small, and often dries up and 
drops off in the autumn, so that only the scale-leaf, v, with the bud (Fig. 1*1 
lt t ) in its axil remains. This bud in the following year developes into a new 
long-branch, and since its leaves lie in a plane at right angles to that of the 
dwarf-branch, their plane coincides with that of the long-branch from which it 
is developed (the grandmother axis). The tendrils no doubt may most correctly 
be regarded as the modified main axis which has been pushed aside by a lateral 
branch. The branches are then sympodia, whose successive shoots bear al- 
ternately 1 and 2 foliage-leaves : thus, on the figure there are portions altogether 
of 5 shoots (I.-V.), the 1-leaved ones are shaded, the 2-leaved ones are white. 
The following facts however are adverse to this theory : (1) the first leaf on an 
axillary bud is then situated 180 from the subtending leaf (e.g. the lowermost 
shaded leaf, Fig. 479, 180 from the lowermost white leaf), whilst the rule in 
the Dicotyledons is that it is placed only about 90 3 to one side. (2) The buds 
(Fig. 479 A'j from which the dwarf-branches develop, must then be accessory 
and sister- buds to the sympoiial shoots, but their first leaves have a different 
relative position to this, which is very peculiar, and a still more remarkable 
fact is that the buds, A', etc. ate similar in structure and present in all the< ; 
thus we only find accessory buds in the cases where no tendrils are opposite 
to the leaves, and the main bud must then be considered to be suppressed. 
(3) The development proves that the tendrils arise on the side of a vigorous 
growing-point of the stem or by its division, and do not develop, as might bo 
expected, from the apex of the shoot. But these relations however, find their 
aualogues and are all capable of explanation, whereas other less natural modes 
of explanation are opposed to them. 

435 species ; especially in the Tropics ; they are rarer in America. In 
N. Am. some Vitii-species and Aiupelopsit quinqnufolia are found. Vitis 
rinifem is supposed to have originated in the districts East and South of the 
Caspian Sea. Wine is obtained from Fi'fis-species, especially V. xinifera, and 
"raisins," (the name " currants," given to a special variety with small, seed- 
less fruits, is derived from Corinth). The specie* of Ampelopsis (Virginian 
Creeper) are cultivated as ornamental plants. 

Order 5. Rhamnaceae. The stamens are placed opposite tie 
petals as in the Ampelidacese (Fig. 482), but the flowers are much 
'more perigynous or entirely eplgynous. The trees and shrubs be- 
longing to this order have simple, mo'st frequently penninerved 
leaves with stipules ; frequently thorny (modified branches). The 
flowers are inconspicuous, sometimes unisexual (Fig. 482), and 
have 5 (-4) sepals, petals, stamens, and generally 3 (2-">) carpels. 
The calyx has valvate aestivation. The petals are very sin nil 
(generally less than the sepals), often spoon-like, hollow, and em- 
bracing the stamens ; a disc covers the inner surface of the thalamus 
or the base of the style in the epigynous flower ; gynceceum simple, 



with one style and one erect ovule in each loculus. The fruit is 
most frequently a drupe. The embryo is large, often green or 
yellow, with endosperm. 

Rhamnus (Buckthorn) has a juicy drupe with 3 (2-4) stones, 
surrounded at the base by the persistent portion of the receptacle ; 
the disc is thin. R. cathartica (common Buckthorn) : dioecious, 
with opposite, serrate leaves. R. frangula (Alder Buckthorn) : 
flowers > , with scattered, entire leaves. Ceanothus (N. Am., with 
richly-flowered inflorescences and a fruit closely resembling that of the 
Euphorbias). Phylica, Pomaderris (Austr., fruit a capsule). Zizyphus, Paliurus, 
Colletia (S. Am.) are thorny shrubs ; C. gpinosa has thorny shoots with small, 
caducous leaves ; the seedling has normal foliage-leaves. Others climb by 
tendrils as in the Ampelidacese, e.g. Gouania. 

FIG. 482. Kliamnus cathartica: A long-styled -flower; pet petals; B short-styled 
cJ-flower; C long-styled ? -flower ; D short-styled $ -flower (after Darwin). 

475 species, 40 genera ; chiefly in temperate and tropical climes. Some are 
medicinal plants, the bark and fruit having purgative properties (the bark of 
Rhamnus frangula and " Cascara Sagrada " from the bark of E. piirshiana are 
officinal). The fruits and seeds of others are edible, for example, the fruits of 
Zizyphus lotus, Z. vulgari*, Z. spin a Christi, etc. Green and yellow dyes are 
obtained from the fruit of R. cathartica, iufectoria and others (Avignon grain). 
Ceanothus-, Rhatnnus- and evergreen Plnjlica-species are ornamental shrubs. 

Family 18. Thymelaeinae. 

Exclusively trees or shrubs with simple, entire, scattered leaves 
without stipules. They have a strongly perigynous, regular, 4- 
inerous flower. The receptacle (often coloured) envelopes a simple 
gynoeceum formed of 1 carpel and with, in most cases, 1 ovule, bear- 
ing on its edge 4 (or 5) petaloid sepals and, but rarely at the same 
time, small, scale-like petals. The corolla is most frequently 
entirely wanting (and hence these plants were formerly reckoned 
among the Monochlamydeas) ; frequently only one of the 2 whorls 
of stamens, which are situated on the inner side of the edge of the 
receptacle, is developed. The fruit is most frequently a \-seeded 


berry or drupe, or a nut which may be falsely berry-like, the 
partly persistent receptacle being fleshy and enveloping it. 

This family appears the most nearly allied to the Frangulinae, especially the 
Khauanaceas, and may be considered as a further development of these in the 
direction of the petaloid development of the receptacle and reduction of the 
corolla and gynoeceum, which in this instance only consists of one carpel. 
Another deviation is that both the whorls of stamens are present, while one 
of these is always wanting in Frauguliua?. They also appear to be related to 
the Lauracete (see page 391). 

Order 1. Thymelaeaceae. The flowers are most frequently $ 
(Fig. 483). The receptacle is high, generally tubular, coloured, 
and bears on its edge the 4- (or 5)-merous calyx, with imbricate 
{estivation. The corolla is wanting or is represented by small 
scales. The stamens are situated on the inside of the receptacle, 
and number 4 + 4 (or 5 + 5) ; stigma capitate. 1 pendulous ovule 

FIG. 483. Daphne mezereum: A flower; B longitudinal section of pistil. 

(Fig. 483 Z>), the radicle pointing upwards. The fruit is most 
frequently a berry. A disc is sometimes developed. Endosperm wanting 
or very slight. 

Daphne (Spurge-Laurel, Fig. 483) has a deciduous receptacle, 
often coloured ; sepals 4 ; petals absent; stamens 4 + 4. Berry. 
Gnidia (corolla) ; Pimelea (2 stamens) ; Tlujmelaia ; Passerina and 

400 species ; chiefly in the warm, sub-tropical zone, especially the Cape and 
Australia. Only Daphne and Thymelaa in Europe. In the fruit and bark of 
some, for example Daphne, pungent, burning and poisonous properties are found. 
The bark of D. mezereum (native and cultivated) and D. laureola is officinal. A 
specially tough bast is found in some species, for example l.ayetta llntearia 
(Lace-tree, Jamaica), which is used in weaving. Some are cultivated in gardens 
as ornamental shrubs, especially species of Daphne. 

Order 2. Elaeagnaceae. Shrubs or trees, which are easily 
recognised by the covering of peltate hairs found upon almost all 
w. B. GO 



parts of the plant, causing- them to assume a silvery or rusty-brown 
appearance. Stipules are absent ; the leaves are simple, most fre- 
quently scattered. Flowers (Fig's. 484, 485) frequently unisexual. 
The sepals are valvate, 2-4 ; the corolla is wanting ; stamens 4 + 4 or 
+ 4. The ovule is erect and the radicle turned downwards (Fig. 486). 
The fruit is a nut, but becomes a, false fruit, being surrounded by 
the persistent receptacle or the lower part of it, and thus assuming 
a berry- or drupe-like appearance (Fig. 486). Endosperm insig- 
nificant. Shepherdia (opposite leaves) has 4 sepals, 4 + 4 stamens, 
as in Daphne. Dioecious. -Elteagnus (Silver-leaf) is ^ > nas 4-6 
sepals, and 4-6 stamens alternating with them. Sippophae is 
dioecious; it has 2 sepals and 4 stamens in the $ -flower (perhaps 
properly speaking 2 + 2 stamens); thorny (stem-structures). 

16 species; especially ornamental shrubs, e.g. Elceagnus arfieiitea, angnsti- 
folia ; Hippopha'i rhamnoides and Shepherdia canadensis. Northern Temp. 

4S4. 485. 486. 

FIGS. 494-486. El&agnus attgustijolia. 

FIG. 4St. Floral diagram. FIG. 485. Longitudinal section through the flower. 
FIG. 436. Longitudinal section through the fruit. 

Order 3 (?). Proteaceae. This order has its chief centre in the dry regions 
of Australia (3^-^, of about 1,000 species), a smaller number in S. Africa dV~A)' 
a few species in S. Am. Trees or shrubs, leaves generally scattered, without 
stipules, and more or less dry, leathery, evergreen, and often of very different 
forms on the same plant (undivided, compound, etc.) The flowers are 
(rarely unisexual), and 4-mermis in the single, petaloid perianth and in the 
staminal whorl ; 1 carpel ; sometimes zygomorphic. The perianth-leaves are 
generally almost free, with valvate aastivation, often leathery. Small scales 
alternating with the perianth are often found at the base of the ovary. The 
stamens generally have extremely short filaments, and are situated opposite, 
Bometimes quite on the tip of the perianth-leaves, in a spoon-like groove. The 
gyuoaceutn is 1-locular, has 1-several ovules, and is often raised on a stalk -like 
internode. The fruit is a follicle or nut. The seeds, most frequently winged, 
have no endosperm. Protea, Maiiylesia, Hahea, Danksia, Grcvillea, etc. 


50 genera ; about 1,000 species. Several species are cultivated in our con- 
servatories for the sake of the flowers, which are beautifully coloured and 
arranged in crowded inflorescences. Protandrous. It is doubtful whether they 
were existent in Europe in the Tertiary Period. The true systematic position 
of the order is doubtful. They are related to the Leguininosaa and Rosiflorse, 
but more closely no doubt to the two preceding orders. 

Family 19. Saxifraginae. 

The flower is generally perfect, regular and polypetalons, 
usually perigynous or epigynous, encyclic and 5-merous ; most fre- 
quently So, P5, A5 + 5 or 5 + and G2-5, but other numbers 
are found, especially 4 ; the flowers are very frequently obdi- 
plostemonous. The calyx is sometimes large and the corolla 
small ; the carpels in some are entirely free, in others more or 
less united. Endosperm is found in the majority. The hypogynous 
forms approach the Cistifloras, the others the following families, especially 
the Rosiflorae. This family is not, upon the whole, so well defined and natural 
as most of the others. The Saxifragaceae proper, approach very near to the 
Rosaceae, especially Spircca, and form a transition to it,. The forms with op- 
posite leaves, as Philadelphia, etc., approach the Myrtifloras, just as the Escal- 
loniaa appear to be closely allied to Bicornes, especially VacciniacecB. Finally 
through Pittosporacetf , they pass over to the Frangulinae. The family ter- 
minates in very reduced forms, on the one hand in the arborescent orders with 
crowded inflorescences, on the other perhaps in the very remarkable order 

Order 1. Crassulaceae. Nearly all are herbs or small shrubs 
with round, succulent branches and scattered, fleshy, often more 
or less round leaves, which are very rarely incised, and never 
have stipules. The flowers are generally 
borne in dichasia or unipared scorpioid e 

cymes, which again may be arranged in 
racemes, umbels, etc. ; they are regular, 
5 , hypogynous or peryginous, and most 
frequently have free sepals and petals 
(gamopetalous corollas with sessile sta- 
mens are found in Cotyledon, Uryophylhim, 
Echeveria, and others); the floral formula 
is Sn, Pn, An + n, Gn, where n may have FlG< 487 ;~ Dia * m of ; ft 

J 6-meroua flower (Sedum IDS- 

very different values, partly depending pnnicum) -. w branch of scor- 

upon the size of the flower (e.g. 4-7 in pioid cyme in the axil of the 

t)rrt.ctGol(^ Q 

Sedum, Fig. 487 ; 6-30 in Sempervivum ; 

4 in Rhodiola, BryophyUum, and Kalanclioe ; 5 in Echeveria, Um- 
bilicus, Cotyledon). The carpels are/ree and are placed opposite the 
petals (Fig. 487). Fruit a syncarp composed of follicles containing 



many, small seeds without endosperm. Outside each carpel is 
found a small, nectariferous scale (Fig. 487). The northern genus, 
Rhodiola, is dioecious. The petal-stamens are wanting in some (Crassula, 
Biilliarda, and others). The floral-leaves are very often displaced upon their 
axillary branches. A multicarpellary gynceceum also occurs. 

Sedum (Stonecrop) is generally 5-merous with 10 stamens ; Sem- 
pervivum tectorum (House-leek), 12-mei-ous, and with 24 stamens. 
The leaves of BnjoplujUum calycinum very readily form buds, and also fre- 
quently exude water from the edges. 

485 species ; especially Temp. (Cape, Europe). Principally used as orna- 
mental plants. 

Order 2. Saxifragaceae. The flowers are 4-5-merous with 2 
(-3) carpels, most frequently: So, P5, A5 + 5 (obdiplostemonous), 
G2. They are regular, ty , polypetalous, hypogynous, perigynous 
or most frequently more or less epigynons (Fig. 488). The carpels 
may be individually quite free, but are more frequently united at 
the base, or the entire portion enclosing the ovules is united into a 
1- or 2-locular ovary, the styles, however, are always free. Fruit 
a capsule with many seeds; endosperm present. They are herbs, 
most frequently with scattered leaves without stipules ; but the 
leaf-base is broad. The inflorescences are most frequently cymose, 
and a displacement of the floral-leaves is frequent (e.g. Clirysos- 
plenium). Some Saxifrar/a-specles, e.g. S. tannenloxa, have irregular flower 
with an oblique plane of symmetry. The petal-stamens in some may be 
wanting : Heuchera, species of Saxifraga and Mitella. The corolla is wanting 
in others. 

Saxifraga (Saxifrage) : S5, Po, A5 + 5, G2 (Fig. 488) ; capsule 
bilocular, opening along the ventral suture between the 2 per- 
sistent styles. S. (jranidnta has small 
tubers at the base of the stem. Chry- 
sosplenium (Golden Saxifrage) : 4 
sepals, no corolla, 4 + 4 stamens; 
1-locular capsule. 


Protandry is most frequently found in 
Sd.rifi'injti, with the stamens successively 
bending towards the gynoecenm; protogyny 
is more rare. In other genera there is 
protogyny without any movement of the 
stamens ; Chrysospleniitm is homogamous. 
About 300 species; mostly in temperate climates. Saxifratiu is especially 
Alpine. S. crassifolia and other species, Hoteia japonica, Tellima, etc., are 
ornamental plants. 

FIG. 483.- Sa.vifi'aga granulata. Longi- 
tudinal section of flower. 



The following genera are allied to the Saxifragacese : - 

1. Parnassia (about 14 species ; P. pahtstris, Grass of Par- 
nassus). The flower is slightly perigynous, and has So, P5, 5 
fertile sepal-stamens, and 5 petal-stamens, which are developed as 
barren staminodes, palmately-lobed, and (3-) 4 carpels united in a 
1-locular ovary with (3-) 4 parietal placentae. Capsule. Protandruus. 
The flower has a slightly oblique plane of symmetry, which is especially 
shown during its development and in the order of sequence in which the 
anthers dehisce : originally they lie closely round the gyuceceuin ; the anthers 
dehisce extrorsely, first the one which is placed opposite the most external 
sepal (the f arrangement is very distinct in the calyx), the filament elongating 
so that the anther lies over the ovary, and this is followed successively by the 
4 others in a zig-zag line ; the filatneuts bend backwards after the pollen is shed 
and the anthers drop off, and the sti^inas are not developed until this is com- 
pleted. The barren stamens are palnmtely divided into an uneven number 
(7, 9, 11) of lobes, tapering from the centre towards the edge, and bearing 
apparently glandular tips; their gland-like appearance is supposed to allure 
flies to visit the flower, or they may act as a kind of fence which compels the 
insects to enter the flower in a certain way, and thus effect pollination; the 
honey is secreted on their inner side, and not by the gland-like tips. 

2. Adoxa moschatellina (Moschatel). This is a perennial, creeping 
herb ; the horizontal rhizome has an unlimited growth, and bears, 
in a scattered arrangement, both foliage-leaves, and white, flesh v 

PIG. 4S9. Portion of Cephalotui follicularis : k pitcher-like leaf with thick corrugated 
edge (m) and lid (I) ; b foliage-leaf of the ordinary form. 

scale-leaves. The aerial stem bears 2 opposite foliage-leaves and a 
capitate inflorescence of 5 flowers, 4 placed laterally (in opposite 
pairs) and 1 terminally. The flower is semi-epigynous, the calyx 



gamosepalous, corolla absent. The stamens are divided to the 
base, so that each filament bears a bilocular anther. The style is 
free, deeply cleft. The terminal flower has 2 bracteoles, 4 sepals, 
4 stamens, cleft to the base, and a 4-locular ovary. The bracts 
of the lateral flowers are displaced on the flower-stalk, as in 
Chrysosplenium, and united with the 2 bracteoles into a kind of 
3-leaved involucre; these flowers have 5 sepals, 5 split stamens 
with 2-locular anthers, and a 5-locular ovary. 1 pendulous ovule 
in each loculus. Fruit a drupe, green-coloured, with 1-5 stones.- - 
This plant, which would perhaps be best placed in a special order, 
has also been classed with the Araliacere and Caprifoliacea?. 

The following are also allied to this order : Escalloniacece (arborescent plants 
with simple, scattered, leathery leaves), Cunoniacea; (arborescent with opposite 
leaves), Cephalotacete (with pitcher-like, insect-catching leaves ; Australia ; 
Fig. 489) and Francoacece, These have respectively 85, 107, 1 and 3 species. 

Order 3. Ribesiaceae (Currants). 5-stamened Saxifragacete 
with epigynous flowers. Moderately sized shrubs with scattered, 


FIGS. 490-492. Ribes rubrum. 

FIG. 490. Floral diagram. FIG. 491. Flower in longitudinal section. 

FIG. 492. Seeds in longitudinal section. 

stalked and palminerved, and generally palmilobed leaves, with a 
large leaf-sheath. The flowers (Figs. 490, 491), most frequently 
borne in racemes, are regular, epigynous, and have often, above the 
ovary, a cup- or bell-shaped, or tubular prolongation of the recep- 
tacle, on which the sepals, petals and stamens are situated; they 
have 5 sepals (often large, coloured), 5 small, free petals, only 5 
stamens (opposite the sepals) and a 2-carpellate gynceceum with a 
unilocular ovary and 2 parietal placentae bearing many ovules. The 
fruit is a berry, whose seeds have a fleshy and juicy outer covering 
(Fig. 492). Iii some species, for example Eiben grossularia, there is found 
an uubranched, or a 3-5-branched spine, very closely resembling the spiny 
leaves of the Herberts, but which, however, are emergences springing from the 
base of the petiole, liibes has two kinds of branches : loug-brai ches and 



dwarf-branches, the latter aloue bearing the flowers. Iiibes (Figs. 490-492). 
The blades of the leaf are folded or rolled together in vernation. 
JR. alpinum is dioecious. 

75 species; especially from the N. Temp, regions (especially N. Am.). The 
receptacle secretes honey on its inner surface. Tbe Gooseberry -flower is 
slightly protandrous, others are homogamous; insect- and self-pollination are 
found. The following are FKUIT BUSHES : K. nig rum (Black Currant), R. rubrum 
(Bed Currant), R. grossularia (Gooseberry), originating in Northern Europe and 
Asia. ORNAMENTAL BUSHES : the North American R. aureunt (Golden Currant) 
and R. saiiyuineiim (Blood-red Currant), etc. 

Order 4. Hydrangeaceae. Shrubs, with simple, opposite leaves, without 
stipules; flowers generally epigynous, 4-5-merous (Fig. 493). Hydrangea (H. 
hortensia, etc.). Shrubs from N. Am. and E. Asia ; corolla often valvate. The 
inflorescence, as in the case of the inflorescence of Viburnum opulus (Guelder 
Eose), has often irregular, large, but barren flowers at 
the circumference, whilst the others are much smaller, 
regular and ; the barren flowers are mostly 
4-merous ; in these cases it is the calyx which is 
large and petaloid, while the other parts of the flower 
are more or less suppressed. The branches of the 
inflorescence appear to be partially devoid of floral- 
leaves, since they are displaced upon the main axis. 
Pltiladelphus; racemes (with terminal flower), sepals 
4 (valvate), petals 4 (twisted), stamens many, and car- 
pels 4 (opposite the petals), forming a 4-locular ovary. 
The numerous stamens (20-30) occur by the splitting 
of the sepal-stamens and are often therefore placed 
in distinct bundles. Fruit a capsule. Ph. coronaria 
(Syringa. Mock Orange-blossom), from S. Eur., is a common ornamental shrub, 
as also is Deutzia (Fig. 493) from N. Am. and E. Asia. The latter has S5, P5, 
A5 + 5, G3. About 70 species. 

Order 5 (?). Pittosporaceae. This order has its home especially in Australia 
(90 species). The flower has S5, P5, A5 (episepalous), G2 (3-5), most frequently 
a unilocular ovary with many ovules in 2 rows, borne on 2 parietal placentae, or 
a bilocular ovary. Some have berries, others capsules. Pittoaparum, Citrio- 
batus, Sollya, Billardiera. 

Order 6. Hamamelidaceas. Flowers more or less epigynous, with S4, PO 
or 4, 4 fertile sepal- stamens, and 4 barren petal-stamens, bilocular ovary with 
1-2 ovules in each loculus. Fruit a capsule. Hamamelis : one species in Japan 
and one in N. Am. Fothergilla. Liquidambar : monoecious ; flowers in capitula 
or spikes ; $ -flowers without perianth, stamens indefinite ; $ -flower : slight 
perianth, 2-locular ovary with rnauy ovules. OFFICINAL : " Styrax-balsam," 
which is obtained by boiling the bark of Liq. orientalis, from Asia Minor. 
Liquidambar and Parrotia are found as fossils in the Upper Oligocene ; 
Hamamelis perhaps in the Chalk. 

Finally two orders with very reduced flowers are included in this family. 
Order 7. Platanaceae. Trees, with large, scattered, palminerved aud'lobed 

FIG. 493. Deutzia crcnata. 
Longitudinal section of 


leaves, and ochreate stipules ; the buds are concealed in a hollow at the base 
of the petiole. The bark falls off in large scales. $ - and $ -flowers (monoecious) 
in crowded, spherical inflorescences which are placed at wide intervals on a 
terminal, thin, and pendulous axis. The flowers have an insignificant calyx 
and corolla; the $ -flower has few stamens ; ? -flower, perigynous, with 4 free 
carpels, 1 pendulum, orthotropous ovule in each. Fruit a nut ; endosperm 
absent. 5 species ; frequently grown in avenues and parks. P. occidentalis 
(N. Am.) ; P. orientalis (W. Asia.). 

Order 8. Podostemaceae. Aquatic plants, especially in swiftly running 
water, with somewhat of an Alga-like, Moss-like, or thalloid appearance ; they 
show themselves in many ways to be adapted to their mode of life and situations 
(having a dorsiventral creeping stem, the flowers sunk in hollows, a formation 
of haptera upon the roots, and thalloid assimilating roots and thalloid stems, 
etc.). Tropical ; 100 species. 

Family 20. Rosiflorae. 

The leaves are scattered, stipulate, or have at least a well de- 
veloped sheath, which is generally prolonged on each side into a 
free portion ("adnate stipules"). The flowers are regular, peri- 
gynous or epigynous. Calyx and corolla 5 ( 4)-?n,erows with the 
usual position. The corolla is always polypetalous. The stamens 
are present in very varying numbers (5-co) and position, but 
always placed in 5- or \Q-merous 'wliorls ; they are frequently 20 in 
3 whorls (10 + 5 + 5 ; see Figs. 494, 502, 505) ; the nearer they are 
placed to the circumference, the longer they are ; they are gene- 
rally incurved in the bud, or even rolled up. The number of the 
carpels is from I-GO ; in most cases all are individually free 
(syncarp), and when they are united it is in every case with the 
ovaries only, whilst the styles remain more or less free (Pomacete, 
species of Spiraea). The seeds have a straight embryo, and usually 
no endosperm. 

The perianth and stamens are most frequently perigynous on the edge of the 
widened receptacle ; its form varies between a flat cupule and a long tube or a 
cup (Figs. 495, 496, 498, 499, 500) ; the carpels are situated on its base or inner 
surface, in some instances on a central conical elongation of the floral axis 
(Fig. 496). The carpels in Pomacece also unite more or less with the hollow 
receptacle, or this grows in and tills up the space between the carpels, so that a 
more or less epigynous flower is formed (Fig. 504). The following numbers of 
stamens occur: 5, 10 (in 1 whorl), 15 (10 + 5), 20 (10 + 5 + 5), 25 (10 + 10 + 5), 
30-50 (in 10-merous whorls) compare the diagrams. The theoretical explan- 
ation of this relation of the 10-merous whorls and their alternation with the 
5-merons whorls is not definitely determined; a splitting of the members of the 
5-merous whorls may be supposed, but the development shows no indication of 
this, and it is not supported in any other way. Several genera have " gynobasic " 
styles, that is, the style springs from the base of the ovary (Fig. 497 A, B). 



The Rosiflora? are on one side closely related to the Saxifragaceae (especially 
through Spircea) from which it is difficult to separate them, and to the Myrti- 
florae ; on the other side they are allied, through the Mimosacaa with the large 
number of stamens, and through the Amygdalaceae with its single carpel, to the 
Leguminosae. The family begins with forms which have many-seeded 
follicles, and passes on the one side to forms with nuts and drupes in perigyuous 
flowers, and on the other side to the Pomacese. 

Order 1. Rosaceae. Herbs or shrubs, generally with com- 
pound leaves and persistent (adnate) stipules, flower perigynous, 
gynoecenm formed of many free (therefore oblique) carpels, syncarps 
with fruitlets of various kinds. The exceptions are noted under 
the genera. 

I. SpIRaiE^! (Fig. 495) has 2-many ovules in each ovary, while 
in the other groups there is generally only 1, and never more than 
2 ovules in each loculus. There are generally 5 cyclic carpels and 
the fruit is 5 follicles, which are not enclosed by the receptacle 

FIG. 491. Diagram of Comarum 

FIG. 495. Flower of Spircea lanceolate. 


The majority are shrubs. Stipules are often wanting. Spirsea 
(Meadow- Sweet). The flowers are generally borne in richly 
flowered inflorescences of various forms. The carpels, in some 
species, unite together and form a simple gynoeceum with free 
styles (an approach to the Pomacese). Closely allied to Spiraea are the 
East Asiatic shrubs : Kerria japonica, which has solitary flowers, in this country 
nearly always double (the fruit a nut), and Rhodotypus kerrioides which has 
opposite leaves, a remarkable feature among the Rosiflorte ; it has a 4-merous 
flower, a well developed disc inside the andrcecium, and a drupe. Closely 
allied also is Gillenia (N. Am.) differing chiefly in the ascending ovules, Spiraea 
having pendulous ovules, and a more tubular receptacle. 

The groups Qiiillajfa and Nnuradece form a transition from Spiraea to 
Pomaceas. In the first group, which contains only trees or shrubs with generally 
simple leaves, the carpels are either free or united (into a capsule) ; in the 
second the receptacle unites with the carpels, which are themselves often united 
together ; in this case, too, the fruit is a capsule. Quill uja (S. Am.) ; E.rochorda 


2. POTENTILLE.E (Figs. 494, 496, 497). The flower has an 
" epicalyx " (Fig. 494 C) alternating with the sepals and formed by 
their stipules which are united in pairs, and hence its leaves are 
often more or less deeply bifid. The receptacle is cupular and 
often quite insignificant. The sepals are valvate in the bud. The 
large number of fruitlets are achenes, borne on a well-developed 
convex portion of the receptacle (the Ranunculeae resemble the Poten- 
tillefe, but Lave no epicalyx, no enlarged receptacle, and spirally-placed stamens). 
Most of the species are herbs with dichasial inflorescences, often 
arranged in racemes. Potentilla (Cinquefoil). The achenes are 
borne on a dry, hairy receptacle ; the style is situated towards the 
apex of the ovary, and is not prolonged after flowering. Herbs 
with digitate, in some, however, pinnate leaves, and generally yellow 
flowers. Comarum (Fig. 494) (Marsh Ciuquefoil) forms, by its 
fleshy-spongy receptacle, a transition to the next genus. Fragaria, 
(Strawberry) (Fig. 496). The receptacle becomes finally fleshy, 

496. A 

FIG. 49C, 497. Fragaria vesca. 

FIG. 49C. Longitudinal section of flower. 

FIG. 4a7. A carpel, entire, and in longitudinal section. 

coloured, and falls off (biologically it is a berry) ; the numerous 
fruitlets (drupes with thin pericarp) have basal styles (Fig. 497) ; 
leaves trifoliate ; long, creeping runners. Geum (Avens) has a 
terminal style which after flowering elongates into a long beak, 
with the apex (after the uppermost part has been thrown off) bent 
back into a hook, thus furnishing a means of distribution for the 

fruits. Leaves pinnate. Dryat comprises 3 Arctic or Alpine species with 
simple leaves and solitary flowers, the calyx and corolla 8-y-merous, the fruit 
resembles that of Geum, but the styles become still longer and feather-like 
(a Hying apparatus). 

3. RUBE.E. liubus (Bramble) has the same form of receptacle 
as the Potentilletv, but no epicalyx; the fruitlets are drupes, not en- 
closed by the persistent calyx. Most frequently shrubs or under- 



shrubs with prickles (emergences), glandular bristles and com- 
pound leaves. In the Raspberry (K. idceus) the fruitlets iinite 
together and detach themselves from the receptacle. 

4. ROSEJC. Rosa ; the receptacle is hollow, ovoid and contracted 
beneath the insertion of the 

calyx (Fig. 498), ultimately 
fleshy and coloured ; it encloses 
a large number of fruitlets 
which are achenes as hard as 
stones (" hip," biologically a 
berry). Shrubs with impari- 
pinuate leaves and adnate 
stipules. The sepals show clearly 
the order of their development (a 
divergence of f), the two outer ones 
oil both sides are lobed, the third 
one on one side only, and the two 
last, whose edges are covered by the others, are not lobed at all. Prickles 
(emergences) are generally present and in some species are placed in regular 
order, being found immediately below each leaf (usually two) although at some- 
what varying heights. 

5. AGRIMONIES. The receptacle is more or less cup- or bell- 
shaped, and almost closed at the mouth ; it is persistent and en- 
velopes the nut-like fruitlets, but is dry, and in some species hard, 
the fruitlets being firmly attached to it. In biological connection 

FIG. 498. Longitudinal sec^on of flower 
of Rosa. 



FIG. 4Q'J, 500. Agrimonia eupatoria. 

FIG. 499. Flo\ver in longitudinal section. 

FIG. 500. Fruit and receptacle in longitudinal section. 

with this the number of the carpels is generally only 1 or 2, and 
the whole becomes a false nut (Fig. 500). Herbs. -Agrimonia 
(Agrimony ; Figs. 499, 500) ; the perianth is 5-merous, stamens 
5-20. The receptacle bears externally, on the upper surface, a 


number of hooked bristles which serve as a means of distribution 
for the 1-2 achenes which are enclosed in it, and hence the 
entire flower finally falls off. The inflorescence is a long upright 
raceme. These bristles are arranged in whorls of 5 and 10, of which the 
uppermost alternate with the sepals. Alchemilla (Ladies-mantle; Fig. 

501) has 8 green perianth-leaves 
in two whorls (some authorities 
consider the four outer as an 
epicalyx, and the flower therefore 
apetalous), and 4 stamens alter- 
nating with the innermost whorl. 
There is only one carpel with a 
basal style and capitate stigma. 
FIG. soi.-Fiower of Aichemiiia in longi- ^he flowers are small and green- 

tudinal section. ' 

ish, the filaments jointed. Ihe 

anthers open by one extrorse cleft. The leaf-sheath entirely 
envelops the stem ; the leaves are palminerved. A. aphanes has 
often only 1-2 stamens. The following genera, with 4-merous flowers 
borne in short spikes or capitula, are allied to this group. Sanguisorba has 
eiitoniophilous, $ -flowers with 4 (-20) stamens, 1 carpel; stigma papillose. 
Poterium ; spike or capitulum, the uppermost flowers are $ , the lowermost $ , 
and some intermediate ones (the order of opening is not always centripetal) ; 
S4, PO, A20-80, G2, the long styles having brush-like stigmas (wind-pollination). 
Leaves imparipinnate. 

POLLINATION. A yellow ring on the inner side of the receptacle, inside the 
stamens, serves as a nectary when any honey is formed ; this, for instance, is 
not the case in Rosa, Agrimonia, Spirtea ulmaria, S. Jilipendula, S. antncu*, etc., 
to which the insects (especially flies and bees) are allured by the quantity of 
pollen. Homogamy and slight protogyny are frequent, in many instances self- 
pollination also is finally possible. Poterium, with the long-haired stigma, is 
wind-pollinated. About 550(1100?) species, especially in northern temperate 
regions. USES. OFFICINAL : the petals of Rosa centifolia and gallica, the fruits 
of the Raspberry (Rubus idtem), the rhizome of Geun nrbanum, the flowers of 
the Koso-tree (Hagenia abyssinica or Brayera anthelmintica). The bark of 
Qitillaja saponaria (Chili) is used as soap and contains saponin. "Attar of 
Hoses" from Rosa dumascrna, centifulia and other species, especially from the 
southern slopes of the Balkans. Many species and varieties of Eoses are 
ORNAMENTAL plants: from S. Europe, Rosa littea (the Yellow Rose), R. tjallica 
(the French Rose) and R. ntbrifolia ; from W. Asia, R. centifolia, of which the 
Moss Roses (R. muscosa audcristata) are varieties, and R. danuiscena ; from India 
and N. Africa, R. moscliata (the Musk Rose) ; from China, R. iudica (Tea Rose) 
etc., besides the native species and the varieties which have been derived 
from them. In addition, Kerria japnnica, species of Potentilla, Rnbm odoratus 
from N. Am., and many species of Spirtea from South-eastern Europe and N. 
Am. ESCULENT : the " hips " of R. niollissima, R. pomi/era, etc. ; the fruits of 



FIG. 502. Diagram of 
Pi'iuius virginiana. 

Rubus-species : Raspberry (R. ida:u$}, Cloudberry (R. chamcemorusj, Blackberry 
(R. fruticoms), etc. ; of Fragaria-species (F. resect, collina, grandijtoia, etc). 

Order 2. Amygdalacese. Trees or shrubs with rosaceous 
flowers ; leaves simple with caducous stipules ; a regular, perigy- 
nous flower, the receptacle being 1 partly 
thrown off bj a circular slit ; sepals 5, 
petals 5, stamens 20-30; gynrjeceum simple, 
formed of 1 carpel (hence oblique, Fig. 502), 
with terminal style and 2 pendulous ovules, 
ripening into a drupe (Fig. 503). The 
leaves are penninerved and frequently have 
glands on the stalks and edges ; thorns 
(modified branches) often occur, i.e. dwarf- 
branches, which, after producing a few 
leaves, terminate their growth in a thorn 
(e.g. Primus spinosa). The vernation of the folia f/e-leaves varies in the 
different genera; in the Almond, Peach, Cherry, and Bird-Cherry they are 
folded; in the Apricot, Plum, Sloe and Bullace, rolled together. In some the 
flowers unfold before the leaves (Amyydalus, Armeniaca). That the gynoeceum 
is formed of 1 carpel is evident in this as in other instances (e.r/. in the 
Leguminosae, which are closely related to this order), from the fact that the 
carpel is oblique, and has only one plane of symmetry, and similarly in the 
fruit tbere ia a longitudinal groove on one side which indicates the ventral 
suture. It is only exceptionally that both ovules are developed. In abnormal 
instances more than 1 carpel is developed. 

A. FRUIT HAIRY: Amygdalus (A. communis, Almond-tree) has 
a dry pulp which is detached irregularly, when ripe, from the 
wrinkled, grooved, ovoid and 
somewhat compressed stone. 
Persica (P. vulgaris, Peach-tree) 
differs from the Almond in hav- 
ing a juicy pulp, not detachable 
from the stone, which is deeply 
grooved and has pits in the 
grooves (Fig. 503). (The name 
of the genus is derived from Persia, 
though it is a native of China). 
Armeniaca (A. vulgaris, Apricot) 
has a hairy, velvety fruit, but 
the stone is smooth and has two FlG - eoa.-Fmit of the Pearh. The pulp is 

cut through so that the stone is visible. 

ribs along one of the edges ; the 

pulp is juicy. (The generic name has been given on the incorrect assumption 

that it was a native of Armenia ; its home is China.) 


B. FRUIT GLABROUS (i.e. without hairs) : Prunus (Plum) has a 
glabrous fruit with bluish bloom ; the stone is compressed, smooth 
or wrinkled. The flowers are borne solitarily or in couples, and 
open before or at the same time as the leaves ; they are borne on 
shoots without foliage-leaves. Cerasus (Cherry) has a glabrous, 
spherical fruit, without bloom, and a spherical stone. The flowers 
are situated in 2 many-flowered umbels or racemes, and open at the 
same time as the leaves or a little before them. Lony-stalked flowers 
in umbel* are found in C. avium (Wild Cherry), C. vuli/arls (the cultivated 
Cherry, from Western Asia) ; racemes at the apex of leaf-bearing branches and 
small spherical fruits are found in C. padus (Bird Cherry), C. virginiana, C. 
laiirocerasus (Cherry-laurel), C. mahuleb. 

POLLINATION. Prtinns spinosa (Sloe, Blackthorn) is protogynous, but the 
stamens are developed before the stigma withprs. Honey is secreted by the 
receptacle. Cerasus jmdun (Bird-Cherry) agrees in some measure with P. spinosa. 
In the flowers of the Plum and Cherry the stamens and stigma are developed 
simultaneously and self pollination seems general ; the stigma, however, overtops 
the inner stamens and thus promotes cross-pollination. DISTRIBUTION. 114 
species in the N. Temp, zone ; few in the warmer regions ; the majority from 
W. Asia. C. vultjaris, from the regions of the Caspian ; Prunus spinoia, 
insititia (Bullace), domesrica (Plum, from the Caucasus, Persia). USES, princi- 
pally as fruit-trees : Cherry, Plum, Apricot, etc. ; " Almonds " are the seeds of 
Amygdalus communi* (W. Mediterranean), "bitter," "sweet," and "shell" 
almonds are from different varieties, the latter being remarkable for the thin, 
brittle stone. In the majority of species and in almost all parts of the plant 
(especially the bark, seed and leaves) is found the glycoside, amyr/dalin, which 
forms prussic acid. Many form gum, and the seeds have fatty oils ("Almond 
oil"). OFFICINAL: the seeds and oil of Ami/fjilalu* communis, and the fruit of 
the Cherry ; in other countries also the leaves of C. laiirocerasus. The stems 
of Cerasu* mahaleb are used for pipes. Ornamental Shrubs: Amygdalus nana, 
Cerasus lanrocerasu*. 

Order 3. Chrysobalanaceae. Tropical Amygdalaceas with zygomorphic 
flower and gyuobasic style. 200 species; especially Am. and Asia. Chrt/soba- 
laiius icaco (Cocoa-plum) is cultivated on account of its fruit (Am.) 

Order 4. Pomacese. Trees and shrubs, most frequently with 
simple leaves and caducous stipules. The flowers (Fig. 505) have 
5 sepals, 5 petals and generally 20 stamens (10 + 5 + 5, or 10 + 10 + 5). 
There are from 1-5 carpels, which unite entirely or to some extent 
with each other, and with the hollow, fleshy receptacle (the flower 
becoming epigynous), (Figs. 505, 506, 507). The carpels are nearly 
always free on the ventral sutures, rarely free at the sides also. 
The whole outer portion of the fruit becomes fleshy, but the por- 
tions of the pericarp surrounding the loculi (endocarp) are most 
frequently formed of sclerenchymatous cells, and are more or less 



firm (the " core "). The nature of the fruit varies, according to 
the thickness and hardness of the endocarp, being either a " berry" 
or a "drupe" (see A and ). When the endocarp is thin and 
parchment-like, the fruit has the characteristics of a berry, each 
of the 5 loculi generally present containing several seeds ; but when 
this is hard the fruit resembles a drupe, only one seed is developed 
in each loculus, and the number of the loculi is reduced to one or 
two. There are nearly always 2 ovules in the loculi of the ovary, 
but in Cydnnia there are a large number in 2 rows. In the genera 

PIG. 505. Floral diagram 
of Msspilus germanic i. 

FIG. 501. Longitudinal ami transverse section through the flowers of A, B Colvneaster ; 
CCydonia; D Malus communis ; E Rapliiolepis ; F Cydonia; G 3fesj>tlus. 

which have stones, only one seed is developed in each stone. The 
genera are distinguished mainly in accordance with the kind of 
fruit and the number of ovules and seeds. 

with thin stone or berrv). 

1. Pyrus and Cydonia; carpels completely embedded in the 
cup-like receptacle, styles always free. Pi/rus : the fruit is 
glabrous, and has only a small calyx, withering or deciduous, 
and a 5-locular ovary with at most 2 ascending ovules in each 



loculus (Fig. 504 D). The large flowers are situated in few- 
flowered umbels or corymbs. P. comnntnis (Pear ; free styles, Fig. 507 ; 
it has the well-kuown pear-shaped fruit ; the core is reduced to several groups of 

FIG. 506. Cydonia vulgaris. Longitudinal section of fruit. 

sclerenchymatous cells embedded in the pulp, the leaf-stalk is as long as the 
blade). Cydtmia (Quince) has a hairy fruit with many seeds in 2 
rows in each loculus of the endocarp (Figs. 504 (7, F ; 506); the 
testa of these seeds is mucilaginous. C. vulgaris, large, terminal 
flowers on lateral branches, 
and large leaf-like, per- 
sistent sepals. 

2. Mains and Amelan- 
chier(Aroiiiu'); carpels free 
on the ventral edge; styles 
united. Mains communis 
(Apple) the fruit is " um- 
bilicate " at the base ; no 
sclerenchymatous cells in 
the pulp; styles united at FIG. 507. Longitudinal section of Pear flower. 

ROSIFLOR^:. 465 

the base (Fig. 504 D) ; leaf-stalk shorter than the blade. Sorbus 
(Mountain-ash) differs only in having a 2-3-locular fruit with 
extremely thin endocarp. Cymose inflorescences in umbellate 
cymes. S. aucuparia has pinnate leaves, S. aria (White-beam) and other 
species have simple leaves. Amelanchier (the Service-tree) has a false divi- 
sional wall springing from the dorsal suture, and more or less projecting into 
each of the loculi of the ovary; Raphiolepis (Fig. 501 E) has racemes and a 
juicy berry; Eriobotrya japonica (Loquat). 

B. CEAT^GEJS. THE ENDOCARP is HARD AND soxr ("drupes," 
generally with several, sometimes, however, with only 1-2 stones, 
rarely one multilocular stone ; only 1 seed in each of the loculi). 
Gratcegus (Hawthorn, May). There are 1-5 stones in the spherical 
or ovoid fruit. The disc, found on the apex of the fruit, inside 
the small, withered calyx, is small (much less than the transverse 
section of the fruit). Shrubs with thorns (branches) and mode- 
rately large flowers borne in corymbs. Mespilus (Medlar) differs 
from the last-named only in having a large disc at the apex of the 
fruit, inside the large, leaf-like sepals, i.e. almost equal to the 
greatest diameter of the fruit. The flowers are solitary and ter- 
minal. Cotoneaster is chiefly distinguished from the others by 
its syncarps, the 2-5 carpels (and stones) being free from one 
another, and only united to the receptacle by a larger or smaller 
portion of their dorsal surface (Figs. 504 A, B). Small shrubs 
with leathery leaves, generally covered with white, felted hairs on 
the lower surface, and with small flowers ; the fruit is red or 

Pear, Apple, Mountain Ash and Hawthorn have prr>togynous flowers which 
secrete honey, aud are conspicuous to ensure insect pollination. 180 species ; 
in the northern temperate regions. Pear and Apple are especially cultivated 
as fruit trees in a number of varieties ; the Paradise Apple (Pynis baccata) ; 
especially in southern countries also the Quince (from N. Persia and the 
Caucasian districts), Medlar and Amelanchier vulgaris. Mains pumila (Caucasus, 
Altai) and M. dasyphylla (Orient, S. Eur.) are regarded as primitive forms of 
the Apple-tree ; M. sylvestris, which grows wild in European forests, appears to 
have been less used. The early Lake-dwellers in Switzerland had the apple- 
tree both wild and cultivated. -The original form of the Pear is supposed to be 
Pyrus achras (Central Asia). Many of tbe species of Cratayiis, some with double 
flowers, and Pyius (Ch<enoiueles) japonica, with brilliant red flowers, are culti- 
vated as ornamental shrubs. OFFICINAL: Quince pips, on account of the 
mucilaginous testa. The fruits contain free organic acids and sugar ; prussic 
acid may be obtained from the seeds. The wood of the Pear-tree is used in 

W. B. H H 


Family 21. Leguminosae. 

The most characteristic feature is, that the gynueceumis \-locular 
and formed of 1 carpel, the ventral suture of which is turned pos- 
teriorly. The fruit, in most instances, is apod (legume), which opens 
generally along both sutures, the two valves twisting more or less 
in opposite directions. In other instances it opens along one suture only, 
or as a pyxidium (Bed Clover), or it is indebisceut, in which case it is more or 
less berry-like (e.g. the Tamarind, Carob-bean), or it is a drupe (e.g. the Ton- 
quin-beau), or a 1-few-seeded nut (e.g. Melilntus), or a lomentum, -which divides 
transversely into as many joints as there are seeds (Omithopus, see Fig. 513). 

The inflorescences belong to the centripetal type (i.e. indefinite) ; 
cymes do not occur. The flowers are zygomorphic, with vertical 
plane of symmetry, seldom regular ; 5-merous with but a few ex- 
ceptions, , and slightly perigynous. The following diagram is 
the most general (Fig. 511) : 5 sepals, with the unpaired sepal 
median and anterior, 5 petals, 5 + 5 stamens, all in alternating 
whorls, 1 carpel. The calyx is most frequently gamosepalous, the 
gyncecenm is narrowed down at the base to a short stalk and, in 
the majority, is more or less bent. The seed is most frequently 
kidney-shaped, with a smooth, hard and shining testa, the hilum 
being very distinct. Endosperm is wanting, or is reduced to a thin 
layer, which is of service when the seed swells during germina- 
tion. The vegetative parts have these features in common, namely, 
the leaves are scattered, stipulate, and almost always compound. 
Peculiar s Jeep -movements and sensitiveness are found in some, chiefly 
in the Mimosas. Many, probably all, Leguminosas have small 
tubercles on their roots which are produced by a kind of bacterium, 
and assist in the assimilation of fi-ee nitrogen. Spontaneous move- 
ments are exhibited by Desmodium gyrans (Telegraph-plant). 

This family is closely allied to the Rosiflorae, with which it agrees in the 
scattered leaves, the presence of stipules, the generally 5-merous and most 
frequently perigynous flowers with eucyclic stamens, and the absence of endo- 
sperm. Amygdalaceiz and Chrysobalanacea:, with solitary carpels, approach on 
one side to the Leguminosae, among which genera with drupes are also found ; 
Mimosacecc, with their many stamens, form a connecting link on the other 
side. In this respect the Mimosa-genus Ajfonsea, and certain Casalpineae and 
Swartzieas, are of special interest in having more than one carpel (syncarp), a 
condition which is sometimes met with abnormally in other Legumiuosaa, as 
will as in AmygJalaceae. About 7,000 species of the Legumicosae are known. 

Order!. Caesalpiniaceae. These are leguminous plants with 
straight embryo and a floicer which is not papilionaceous and has not 



the same aestivation (Figs. 508-510); but in reality there is not a 
single characteristic which, absolutely distinguishes them from the 
Papilionacege. The majority are aborescent ; the leaves as a rule 
are pinnate or bi-pinnate. The flower is 5-merous, most frequently 
perigynous and slightly zygomorphic ; the calyx is free or 
gamosepalous, the corolla polypetalous with ascending imbricate 
aestivation (i.e. the two lowest petals envelop the lateral ones, 
and these again the posterior; Fig. 508) ; 10 free stamens] fruit 

Cassia (Figs. 508-510) is the largest genus (about 200 species) ; 
it has an almost hypogynous, zygomorphic flower with 5 free 
sepals and petals ; of the 10 stamens the 3 posterior are generally 
barren, the others are of very unequal length and open at the 
apex by pores (Fig. 509). In some (the Senna group) the fruit is 

FIG. 508. Floral diagram. 

FIGS. 508-510. Cassia floribunda. 
FIG. 503. Flower. FIG. 510. The same in Ion?, sect. 

a flat, short, thin, dehiscing pod ; in others (Cathartocarpus) it is 
round, long, woody or fleshy, indehiscent, and divided internally 
by more or less fleshy transverse walls into as many cells as there 
are seeds. The following also have DEHISCENT FRUITS : Bauhinia 
(often lianes, tropical climbers with tendrils [stem-structures] and 
anomalous stems), Copaifera, Ha-.matnxylon (whose pod does not 
dehisce along the suture, but laterally), Cercis (simple leaves; the 
corolla resembles that of the Papilionacea?, but the posterior petal 
is the smallest, and is enveloped by the 2 lateral ones, which are 
enveloped in their turn by the 2 anterior). FKUIT INDEHISCENT: 
Tamarindus indica ; the pod is almost round, often a little 
abstricted between the seeds ; the wall is formed by a thin, 
brittle external layer, enclosing an acid pulp ; well-developed 


septa are present, between the seeds ; the most internal layer is 
parchment-like. Calyx 4-merous by the coalescence of 2 sepals. 
Only 3 fertile stamens. Ceratonia siliqua (Carob-bean, Locusts) ; 
the pod is long, compressed, with thick sutures, and has a wall, 
the central part of which is more or less leathery, fleshy and 
sweet ; there are transverse septa between the seeds, as in the 
Tamarind. Embryo greenish in endosperm. The flower is without 
a corolla, 5 stamens. Pterogyne (winged fruit), etc. KRAMERIE^; 
with Kraineria is an anomalous group. 

DISTRIBUTION. 80 genera, with 740 species; almost exclusively in the Tropics. 
The Carob-tree and Cercis grow in the Mediterranean basin. The largest and 
most widely distributed genus is Cassia, which is found as trees, shrubs, and 
weeds in all tropical countries. The order has many important uses to man- 
kind. MEDICINAL : the leaves and pods of Cassia acuti folia and aiip-ustifolia 
(officinal, Senna-leaves), the fruit-pulp of the Cam'a-sub-genus, Cathartocarpus. 
Bhatany root from Kraineria triandr.i (Peru, officinal). Balaam is extracted 
from a number of Co^ai/era-species (Balsam of Copaiba) from S. Am. (offi- 
cinal), and from Hymentea (Copal balsam), Tracltylobinm and others. Edible 
fruits are obtained especially from the Carob-tree (from the East) and the 
Tamarind (officinal). The heart-wood of several species of Ccusalpinia, such 
as C. bradliensis (the Pernambuco-tree), echinata (Bed-tree), and tappan, 
yield dyes ; Hccmatoxylon (H. campechinmcm, Logwood), Copaifera Iracteata 
(Amarant-tree). Timber is obtained from many (Melanoxylon and others). 
In Europe they are of little importance as ornamental plants, these being con- 
fined principally to the species of Gleditschia (G. triacantha, from N. Am.) 
and Cercis (the Judas-tree, C. siliqua strum, S. Eur.), which are cultivated in 
gardens; but in tropical gardens beautiful flowering species, e.g. of Cassia, 
Poinciana, Brownea, are found, and the most beautiful of all ornamental plants, 
the Indian Ainlierstia nobilis. 

Order 2. Papilionaceee. The flower (Figs. 511, 512) is 
strongly zygomorphic and somewhat perigynous (Fig. 512 U ; most 
frequently more on one side than the other). The calyx is 

gamosepalous and persistent. The polypetal- 
ous corolla has descending imbricate estiva- 
tion, the posterior, large leaf, the standard 
(Figs. 511 /; 512 B', e), covering in the bud 
the two lateral ones, the icings (Figs. 511 v; 
'^12 B', a), which again cover the two an- 
terior ; these are united in the form of a boat, 
the keel (k and c) ; the wings and the two 
FIG. 5ii. Diagram of petals of the keel are very unsym metrical. 

Faba vulgarls: / the That the kee] ^ f orme d of two petals is 

standard ; v the wings ; 

k the keel. seen by its position (in front of one sepal) 


and by the two often more or less free claws. The 10 (5 + 5) 
stamens (monadelphous) are either all u