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CAMBRIDGE BIOLOGICAL SERIES.
GENERAL EDITOR ARTHUR E. SHIPLEY, M.A., F.R.S.
:

FELLOW AND TUTOR OF CHRIST'S COLLEGE, CAMBRIDGE.

THE
BRITISH FRESHWATER
ALG^E
 ILonfcon: C. J. CLAY AND SONS,
CAMBRIDGE UNIVERSITY PRESS WAREHOUSE,
AVE MARIA LANE,
: 50, WELLINGTON STREET.

ILctpjtg: F. A. BROCKHAUS.
JJeto Jforfe: THE MACMILLAN COMPANY.
Eombao anS (Calcutta: MACMILLAN AND CO., LTD.

[All Rights reserved.]

 *
<-,.

Plankton from Lough Neagh, Ireland (

x 100).

I'lankton from Loch Buar, Sutherland (xlOO).

 A TREATISE
ON THE

BRITISH FRESHWATER
ALG.E

BY

G. S. WEST, M.A., A.R.C.S., F.L.S.,

PROFESSOR OF NATURAL HISTORY AT THE ROYAL AGRICULTURAL COLLEGE,
CIRENCESTER FORMERLY SCHOLAR AND HUTCH1NSON RESEARCH
;

STUDENT OF ST JOHN'S COLLEGE, CAMBRIDGE.

CAMBRIDGE
AT THE UNIVERSITY PRESS
1904
 Cambridge :

PRINTED BY J. AND C. F. CLAY,

AT THE UNIVERSITY PRESS.

 PREFACE.
a
object of the present volume is to give the student
T'HE
concise account of the structure, habits and life-histories of
Freshwater Alga?, and also to enable him to place within the
prescribed limits of a genus any Alga he may find in the fresh
waters of the British Islands.
Although it may seem incongruous to treat of freshwater Algse
apart from marine ones, there are many excuses for the production
of a special on the freshwater, forms. Few genera of
treatise
and fewer
still species, exist both
in salt and fresh water,
Algae,
and the vast majority of marine Alga^ are very different in nature
from those inhabiting fresh water.
The need for a book of this kind is
very great, owing to the
rapid strides made in the investigation of this class of plants

during the last twenty years. To identify even many of the

commonest of freshwater Alga? one has at present to be fully
conversant with most of the recent phycological literature, and
I have endeavoured in this volume to give a general account of
those facts concerning the British species which will be of most
assistance to a diligent student.

Many facts and suggestions concerning the life-histories, de-

velopment, and relationships of freshwater Algae are here brought

forward for the first time, and with few exceptions the figures are
original, pains having been taken to state
as far as possible the
localities from which the specimens were collected. The figures
are careful and accurate drawings to scale, and they are in no wny

diagrammatic. A
few stages in the life-histories of various Alg;c.
and certain figures showing structural peculiarities, have been
copied from the original drawings of other authors,
but in each
case this has been specially mentioned.
vi Preface

The magnifications given under the figures are far from uni-
form, but this is no great disadvantage, as a knowledge of the
'

relative sizes of these plants is very soon acquired, and size is

'

of no generic and little specific value.

Accurate measurements are given of the most abundant and
widely distributed species.
It was originally my intention to add a chapter on the Pcri-
diniea:>, but after further consideration I have for two reasons

excluded them. Lack of space would have compelled me to have

given a veiy indifferent account of them, and I also prefer to
regard them as a group quite distinct from the Algae. Most
zoologists claim these organisms as Dinoflagellates, and the Peri-
dinieas of this country are sufficiently numerous and important
to require separate treatment.
Similar remarks apply to the Characese. They are best re-
garded as distinct from the Alga3, as their vegetative organs
exhibit far more differentiation than the vegetative structures of

any of the freshwater Algse, and their sexual reproduction is of a

distinctly higher type.

In some instances I have quoted freely from previous publi-
cations of my own, sometimes with slight alterations.
The work was undertaken at the request of Mr A. E. Shipley,
to whom I tender my best thanks for assistance and advice during
itspublication. I have also to thank Mr Edwin Wilson for the
care he has expended in reproducing the drawings.
I take this opportunity to remark that a good systematic text-

book of British Freshwater Alga?, with descriptions of all the

known species, remarks on their affinities, and if possible, with
figures, is at the present time very urgently needed. There is, in
fact,no single book, or set of accessible books, by means of which a
student can hope to accurately identify one-third of the freshwater
Algae he may find in a single day's ramble through a reasonably
productive part of the country. I venture to state that this is
the only branch of systematic botany in which such a state of
affairs exists.

The reason for it is not far to seek. It is one result of the

general neglect of systematic botany (especially Cryptogamic) in
the botanical laboratories of this country. Every encouragement
 Preface vii

has been given, and given, to students to take up physiological

is

botany or to investigate the morphology of such plants as exhibit

sufficiently striking peculiarities to arrest the attention even of
a casual observer; but what encouragement is given to a student
who wishes to take up the systematics of any group of plants ?
The answer is found in the Bibliography of every work on
' '

systematic botany.
Although the work of the systematist is indispensable to a
laboratory worthy of the name, there is undoubtedly in many
quarters a lack of appreciation of systematic work, because it is
at the same time the most laborious and the most vexatious of

any form of biological investigation. It is becoming more and

more difficult every day to conduct systematic investigation away
from the special libraries indispensable to every well-equipped
laboratory, and unless a slight encouragement is given to the
laboratory student to take up some branch of systematic botany
thisdepartment of botanical science will be left largely in the
hands of the foreigner.
One cannot emphasize much
the importance of a sound
too

knowledge of the geographical distribution of some of the more

lowly types of Cryptogams, particularly of the Desmidiaceas.
Such a knowledge, which can only be acquired by the patient
labours of the systematist, will throw much light on one of the
most interesting of problems concerned with the later phases
all

of the earth's history, namely, the land-connections of previous

periods.
The frontispiece consists of a reproduction of two photo-
micrographs to show some of the characters of the freshwater
plankton. One is a photograph of some material collected from
Loch Ruar, Sutherland, by Mr J. Murray, of the Scottish Lake
Survey (Pullar Trust). The other represents plankton of a
somewhat different nature from Lough Neagh, Ireland.

G. S. WEST.

ClRENCESTER,
April 7th, 1904.
 TABLE OF CONTENTS.

INTRODUCTION.

PAGE
Books relating to British Freshwater Algae 1

Cultivation of Algae .....

Occurrence, collection, and preservation of Freshwater Algae . .

9
3

ALG^E.

Algfe what they are, and the six classes into which they are divided
:
10
Structure of Freshwater Algae 11
Vegetative multiplication
Asexual reproduction .........
.....
13
14
Sexual reproduction
Polymorphism
Phylogeny and
..........
........
classification
. 15
18
21

Class 1. RHODOPHYCE.E, p. 34.

Order I. NEMALIONACE^E .36

Family 1. Helminthocladieae 36
Batraohospermum, Chantransia, Thorea
Family 2. Lemaneacese 40
Lemanea, Sacheria
Order II. CRYPTONEMIACE.E ... 43

Family 1. Squamariaceae 43
Hildenbrandtia
x Contents
PAGE

Class 2. PH^EOPHYCE^E, p. 44.

Order I. SYNGENETICLE . ... 44

Family Hydruraceae
1. ... 45

Family
Hydrurus
-2.
Chrysomonadinacese .... 46

Synura, Syncrypta, Uroglena

Family 3. Dinobryacese 47

Dinobryon
Family 4. Phseocapsacese 48
Phseococcus, Pbseospheera, Stichogloea

Class 3. CHLOROPHYCE^E, p. 50.

Order CEDOGONIALES 57

....
I.

Family 1. (Edogoniacese 57
(Edogoniurn, Bulbochsete

Order II. CH^TOPHORALES 66

Family 1. Coleochsetacese . 67
Coleochsete

Family 2. Herposteiraceae ... . 70

Herposteiron
3. Ulotrichaceas 73
Family .

Ulothrix, Hormospora, Geminella,

Gloeotila, Radio-
tilum, Stichococcus, Uronema, Binuclearia

Family 4. Cylindrocapsacese . . 81

Cyliudrocapsa
5. Chsetophoraceae 83
Family . .

Cha'tophora, Myxonema, Draparnaldia, Pseudochsete,

Thanuiiochsete

Family 6. Microthamniacese
Microthamnion, Gongrosira, Leptosira
Family 7. Trentepoliliacese . .
93

Trentepohlia
Order III. ULVALES . 95

Family 1. Ulvaceae 95

Moiiostroma, Eiiteromorpha
Order IV. SCHIZOGONIALES .... 98

Family 1. Prasiolaceae
Prasiola
... 98
 Content* \\

Order V. MICROSPORALES '. ... .100

I'Ai.F.

Family 1. Microsporaceae . . loo

Microspora
Order VI. CLADOPHORALES 101

Family 1. Cladophoraceae . . Kci

Chsetomorpha, Rhizoclonium, Cladophora, Chsetonella

Family 2. Pithophoraceae ... . . 106
Pithophora
Family 3. Sphaeropleaceae . ... Mty

Sphseroplea
SIPHONED
Order VII.

Family I. Vaucheriaceae
Vaucheria
. .

....... 108
i<>'.

CONJUGATE .114
Order VIII.

Family 1. Zygnemaceae ........

. .

.117
116

Sub-family I. MESOCARPE^E . . .

Mougeotia, Gonatonema
Sub-family II. ZYGNEMEiE ...... 123

Family 2. Desmidiaceae
SACCODERM^E
.......
Debarya, Zygnema, Spirogyra, Choaspis

...... 135

152
Sub-family
Tribe 1.
I.

Gonatozygce
Gonatozygon, Genicularia
....... 152

Tribe 2. Spirotceniece . . . . . . . I "> t

Spirotsenia, Mesotsenium, Cylindrocystis, Netriurn

Sub-family
Tribe 3.
II. PLACODERM^E
Peniece
......
...... 156
1">7

Penium
Tribe 4. Closteriece . . . . . . . 157

Tribe
Roya, Clo.steriutn
5. Cosmariece .....
Docidium, Pleurotsenium, Tetiiu-morus, Eiiastrum,
. 161

Micrasterias, Cosmarium, Xanthidium, Arthro-

desmus,Staurastrum,Cosmocladium,Oocardium,
Spluerozosma, Onychonema, Spoiidylosium, Hy.-i-

lotheca, Desmidium, Gymno?.vu;i

Order IX. PROTOCOCCOIDE.E .... 178
180
Family 1. Chaetopeltideae . . .

Chsetopeltis, Chaatosphaeridium, ( 'omu-liii'to, Poly-

chsetophora
xii Content*
PAGE

Family 2. Volvocacese 184

Sub-family I. CHLAMYDOMONADE/E 186

Carteria, Chlamydomonas, Chlorogonium, Sphserella
Sub-family II. PHACOTE^E 189
Phacotus

Sub-family III. VOLVOCE^E 190

Goniurn, Stephanosphsera, Pandorina, Eudorina, Volvox

Family 3. Endosphaeracese . . . 197

Family 4. Characieae
Characium
...
Chlorochytrium, Centrosphoera, Phyllobium
199

5. Pleurococcaceae 201
Family . . .

Pleurococcus, Trochiscia, Radiococcus, Protoderrna, Hor-

motila, Urococcus

6. Hydrodictyaceae 206
Family
....
. .

I. HYDRODICTYE^E 207
Sub-family
Hydrodictyon
209
Sub-family II. PEDIASTRE.E . .

Pediastrum, Euastropsis
7. Protococcaceae Autosporacese) . 212
Family (or

I. CCELASTRE^ . . . 213
Sub-family
Coelastrum, Sorastrum

II. CRUCIGENIE^E 215

Sub-family .

Crucigenia, Tetrastrum

Sub-family III. SELENASTRE.E .... 217

Dactylococcus, Scenedesmus, Dimorphococcus, Ankistro-

desmus, Closteriopsis, Actinastrum, Selenastruni,
Kirchneriella

Subfamily IV. OOCYSTIDE/E .... 226

Oocystis, Nephrocytium, Eremosphsera, Palmellococcus,

Chlorella

231
Sub-family V. TETRAE'DRIE^E . .

Tetracdron, Cerasterias

PHYTHELIE.-E 232
Subfamily VI. . . .

Golenkinia, Eichteriella, Lagerheimia, Chodatella

DiCTYOsrHjERiEJE 235
Sub-family VII. . .

Dictyosphoerium, Dictyocystis, Tetracoccus, Botryococ-

cus, Ineffigiata
 Content* xiii

Family 8.

Sub-family
Palmellaceae
I.
........
PALMELLE^E ......
PAGE
239
:j|o

Sub-family II. ......

Palmella, Palmodactylon, Schizochlamys, Sphserocystis
TETRASPORE.E -2\:>,

Tetraspora, Apiocystia

Sub-family III. ......

GLCEOCYSTIDE^E
Glceocystis, Dactylothece, Palmodictyon, Botrydina
-i \ \

Class 4. HETEROKONT^E, p. 248.

Order I. CONFERVALES
Chlorotheciaceae
........
....... 249
250
Family 1.

Oodesmus
Family 2. .......
Stipitococcus, Characiopsis, Mischococcus,
Tribonemaceae 253

Family 3. Botrydiacese
Botrydium
........
Chlorobotrys, Ophiocytium, Tribonema, Bumilleria
258

Class 5. BACILLARIE^E, p. 260.

Order I.

Sub-order
CENTRIC^
1. DISCOIDE.E
........
........ 273
274

Family 1. Melosiracese . . . -74

Family
Melosira
2. Coscinodiscacese ....... 276

Sub-order 2.

Rhizosoleniacese
.......
Cyclotella, Stephanodiscus, Coscinodiscus
SOLENOIDE.E ^77

Family 1. .

Rhizosolenia, Cylindrotheca

Order II. PENNATJ5 .... 279

Sub-order 1. FRAGILARIOIDE^ . . 280

1. Tabellariaceae . 281
Family . .

Tetracyclus, Tabellaria, Diatomella, Denticula

Family 2. Meridionacese
Meridion
Family 3. Diatomacese .

Diatoma
Family 4. Fragilariaceae
Fragilaria, Synedra, Asterionella

Family 5. Eunotiaceae .

Ceratoneis, Eunotia
xiv Contents
PAGE

Sub-order 2. ACHNANTHOIDE^E . . 289

Family 1. Achnanthaceae
Achnanthes

Family 2. Cocconeidacese
Cocconeis

Sub-order 3. NAVICULOIDE.E .
291

Family 1. Naviculacese .

Navicula, Stauroneis, Vanheurckia, Ampbipleura, Gyro-

sigtna, Amphiprora, Mastogloia
~
29
Family 2. Gomphonemacese .

Gomphonema, Rboicosphenia
Family 3. Cocconemacese
Cocconema, Amphora, Epithemia
Sub-order 4. NITZSCHIOIDE^E .
301

Family 1. Nitzschiacese
Bacillaria, Nitzschia, Hantzschia

Sub-order 5. SURIRELLOIDE^ . 303

303
Family 1. Surirellaceae . .

Cymatopleura, Surirella, Campylodiscus

Class 6. MYXOPHYCE^, p. 306.

Sub-class 1. GLAUCOCYSTIDE.E . . 316

Family 1.Glaucocystacese 317

Glaucocystis, Chroothece

...
Sub-class

Order
2.

I.

Sub-order
ARCHIPLASTIDE.E
HORMOGONE/E
1. PSILONEMATE.E
... . .
.318
.319
317

Family 1. Stigonemaceae . 319

Stigonema, Hapalosiphon

Family 2. Scytonemacese . 322

Microchrete, Scytonema, Tolypothrix, Desmonema, Diplo-
colon

Family 3. Nostocaceas 324

Nostoc, Anabtcna, Aplianizomenon, Nodularia, Cylindro-
spermum
 Contents xv
PAGE
Family 4. Oscillatoriacese 329

Sub-family I. VAGINAKIE^ 330

Schizothrix, Dasyglcea, Microcoleus

Sub-family II. LYNGB^E^; 332

Plectonema, Symploca, Lyngbya, Phormidium, Oscil-

latoria, Arthrospira, Spirulina

Sub-order 2. TRICHOPHORE.F. . 337

Family 1. Rivulariacese 337

Amphithrix, Calothrix, Dichothrix, Rivularia, Glceo-
trichia

Family 2. Camptotrichacese 341

Amrnatoidea

Order II. COCCOGONEJ3 . . 342

Family Chamaesiphoniaceae
1. 342
Chamsesiphon
Family 2. Chroococcaceae 343

Sub-family I. CHROOCYSTE^E 344

Gloeocheete

Sub-family II. CHROOCOCCE.E 345

Gloeothece, Aphanothece, Synechococcus, Dactylococ-
co})sis, ^lei'ismopedia, Tetrapedia, Coelosphperium,
Gomphosphoeria, Microcystis, Glceocapsa, Aphano-
capsa, Porphyridium, Chroococcus

INDEX 3">3

ERRATA.

Page 3, line 19, for Glaucocystids read Glaucocystidea\

,, 14, last line, after Zyguemaeere insert a semicolon.

,, 165, line 12, for E. ciimatum Ehrenb. rend K. aiifatum llsilfs.

304, line 1, for Cymatopleura Turpin 1827, read Cymatopleura W. Sm.,

1851.

,, 358 .(Index), for Dabarya read Debarya.

xiv Contents

PAGE
Sub-order 2. ACHNANTHOIDE^E 289

Family Achnanthaceae
1. 289
Achnanthes

Family 2. Cocconeidaceae 290

Cocconeis

Sub-oi-der 3. NAVICULOIDE^E 291

Family 1. Naviculaceae 291

Navicula, Stauroueis, Vanheurckia, Amphipleura, Gyro-
sigma, Amphiprora, Mastogloia
Family 2. Gomphonemacese 297
Gomphonema, Ehoicosphenia
Family 3. Cocconemaceae 298
Cocconema, Amphora, Epithemia
Sub-order 4. NITZSCHIOIDE.E 301

Family 1. Nitzschiaceae 301

Bacillaria, Nitzschia, Hantzschia

Sub-order 5. SURIRELLOIDE.E ...... 303

Family Surirellaceae
1. 303
Cymatopleura, Surirella, Campylodiscus

Class 6. MYXOPHYCE.E, p. 306.

Sub-class 1. GLAUCOCYSTIDE.E .
316
 Contents xv
PAGE
Family 4. Oscillatoriacese 329

Sub-family I. VAGINARIE^E 330

Schizothrix, Dasyglcea, Microcoleus

Sub-family T [.
LYNGBY^E 332
Plectonema, Symploca, Lyngbya, Phormidium, Oscil-
latoria, Arthrospira, Spirulina

Sub-order 2. TRICHOPHORE.E . .337

Family 1. Rivulariaceae 337
Amphithrix, Calothrix, Dichothrix, Rivularia, Gloeo-
trichia

Family 2. Camptotrichacese 341

Ammatoidea
Order II. COCCOGONE.E 342
Family 1. Chamaesiphoniacese 342
Chamaesiplion
Chroococcaceae
..... 343
Family 2.

Sub-family I. CHROOCYSTE^E 344

Gkeochtete

Sub-family II. CHROOCOCCE.E 345

Gloeothece, Aphanothece, Synechococcus, Dactylococ-
copsis, Merismopedia, Tetrapedia, Coelospheeriurn,
Gomphosphseria, Microcystis, Gloeocapsa, Aphano-
capsa, Porphyridium, Chroococcus

INDEX . .
353

Plankton from Lough Neagh and Loch Ruar .... Frontispiece

 INTRODUCTION.
ONE of the earliest attempts to bring together all that was
then known concerning British Freshwater Algas was Dillwyn's
British Conferva,' which appeared in 1809, and hardly any further
'

advance was made in Britain until the appearance in 1845 of

Hassal's History of British Freshwater Algae.'
'
About this time
two very important works were commenced on the continent, one
being Kiitzing's Tabulae Phycologicae,' the first part of which
'

appeared in 1846, and the other Rabenhorst's 'Flora Europsea '

Algarum,' issued from 1864-68. Ralfs' British Desmids appeared

'

in 1848, and for the next thirty years Henfrey, Hicks and Archer
were almost the sole contributors to the literature of British Fresh-
water Alga?, the publications of Archer being very numerous and
most valuable.
'

From 1882-84 Cooke issued his British Freshwater Alga?

'

and in 1887 Wolle's 'Freshwater Algae of the United States'

appeared. Since the publication of these two
books more actual
work has been accomplished in the investigation of freshwater
Algge than at any previous period, particularly by continental
investigators, amongst whom may be
mentioned Wille, Wittrock,
Nordstedt, Bornet, Thuret, Lagerheim, Klebs, Hansgirg, Schmidle,
Chodat, Borge, Boergesen, Lemmermann, and others. In Britain,
Marquand, Bennett, and Roy have done systematic work in certain
districts, and, in conjunction with my father, I have myself spent
much time in the investigation of the Algae of many parts of the
British Islands. During this later period of approximately twenty
in the life-histories
years many new phases have been discovered
of Algae, and much has been found out with regard to their habits
and mode of life. In addition, a great deal has been accomplished
in clearing up the synonymy of these plants, so that taking into
consideration all these circumstances, it is now doubtful if thirty
w. A.
2 Introduction

per cent, of the British freshwater Alga?, could be identified with

certainty from Cooke's book and Wolle's American publication
;

would be of little The relationships and affinities

or no assistance.
of the Algse described in these publications, and in many cases
their life-histories also, are now much better known. This has
resulted in great changes in their classification.
At the time Cooke's book was published it was expected by
many that during the next few years the groups of the Protococcacese
and Chroococcacese would have disappeared, and statements were
made by certain authors advocating great polymorphism in Algse,
evidence being brought forward to prove that many of the more
lowly forms were obviously stages in the development of higher
forms. I have elsewhere pointed out that these statements were
1

based upon very inadequate observations and that more evidence

is yet required concerning the life-histories of some of the lowly

types before any definite statements can be formulated as to their

systematic position. Again, because a few observers have at
different times suggested and attempted to show that these lowly

types are only developmental stages, that constitutes no reason

why, when their life-histories are unknown, they should be neglected
and left out of consideration. Yet, that is largely the case in Wolle's
'
Freshwater Algse of the United States.'
Blackmail 2 in advocating the primitive nature, of the genus
"
Chlamydomonas, also remarks that this specific constancy of the
most primitive type is in strong opposition to the idea of wide
polymorphism brought forward by Hansgirg, Chodat, and Borzi,
which associates different genera, and even members of different
families, in the life-history of one individual." No one would
doubt for a moment the existence of considerable polymorphism
in most groups of the Algse, but it is certainly on a more reason-
able scale than was at one time advocated.
The classification I have adopted is thoroughly explained in
the next chapter and the reasons for so arranging these plants are
stated in full.

With regardto the terminology, I have for the most part

followed that used by Vines in his Text-book of Botany 3
'

.'

1
G. S. West, 'Algafi. of Cambridgeshire,' Journ. Bot. 1899, pp. 5253.
2
F. F. Blackman, 'The Primitive Algae and the Flagellata,' Ann. Bot. xiv, 1900,
p. 660.
3
S. H. Vines, 'A Students' Text-book of Botany,' London, 1895.
 Introduction 3

Exception may be taken to the rejection of Cohn's group of

the Schizophyta, which was instituted to include the SchizomycH 's >

(or Bacteria) and the Schizophycese (or Blue-green Algae). These

two series of plants undoubtedly present a similarity in their
method of multiplication by simple cell-fission, but it must be
remembered that most unicellular and colonial Alga> habitually
multiply in this manner, and although the Bacteria stand near in
this respect to some of the less differentiated
Blue-green Algrc,
there are many reasons for not including the two series of organisms
in the same group. The Blue-green Algae
possess chlorophyll and
phycocyanin disposed within the cells in the manner of a primitive
chromatophore, and are thus capable of carbon-assimilation whereas ;

in the Bacteria this is not the case. A ciliated motile condition

isonly known to occur in about two blue-green organisms, whereas
that is the normal condition in most of the Bacteria and the ;

spore-formation, with very few exceptions, is quite different in the

two groups. The Myxophycese are also of a much higher type of
organization than the Bacteria, possessing a primitive nucleus
(which in the Glaucocystidae has become a true cell-nucleus) and a
cell-wall composed partly (and in the case of certain cells, entirely)
of cellulose. Moreover, the habits and mode of life qf most of them
are totally different from those exhibited by the Bacteria.
In this volume the Blue-green AlgaB are placed in the class
Myxophyceas (Stizenberger, I860) which is an earlier name than
1
,

Phycochromophyceae (Rabenhorst, 1864) or Cyanophyceae (Sachs,

1874), the limits of which were carefully and exactly made out by
Stizenberger. It is the name under which the Blue-green Alga 1

have been placed by systematists for many years past, but for
some unaccountable reason it has not up to the present been
even mentioned in general text-books on botany.

OCCURRENCE, COLLECTION, AND PRESERVATION OF

FRESHWATER ALG.-E.

Algas are universal in their occurrence, no moist situation being

without some type of Alga. They are found on damp earth, rocks,
walls, palings, tree-trunks, in rain-tubs, etc. they are met with in
;

all kinds of running water, from the torrent, waterfall and cataract

1
Stizenberger in Rabenhorst's Algen Sachsens systematise!* geordnet, 18(JO, p. 17.

12
4 Introduction

to the slowest river. Theyare most abundant, however, in still

waters, occurring in quantity in pools, ditches and lakes.
They occur either floating at the surface, being simply immersed
in the water, or attached tosubmerged stones, or to larger aquatic
plants as epiphytes, etc. The larger Algse are often conspicuous
as green slimy masses forming a surface coat to ponds, ditches, etc.,
or attached as large green masses to the rocks and stones of rivers.
Rocks over which the water is constantly dripping possess at times
quite a characteristic Alga-flora, and many of the more uncommon
Alga? are found in such situations. If such rocks present vertical
wet faces, they are often covered with thick leathery patches, or
gelatinous masses, which exhibit a great variety of colour. This
material is always of interest and usually consists of a mixture of
plants belonging to the blue-green Alga? or Myxophycea?.
Some Algae have acquired a symbiotic relationship with other
plants, and even with animals. One species of Anabcena lives
symbiotically with the aquatic Lycopod Azolla, and another with
the Hepatic Blasia, and some Alga? belonging to the genus
Chlorella are connected symbiotically with such animals as
Hydra and certain small species of Amoeba, Paramecium,
vlridis

Ophrydiwn, etc. Again, there is the Lichen, which is a compound

organism consisting of a Fungus associated symbiotically with
Alga? of the genera Plearococcus, Trentepohlia, Cephaleuros, Nostoc,
Stigonema, Scytonema, etc. The genus Foreliella has a symbiotic
relationship to the freshwater mussel (Anodonta) and several other
Algse are similarly related to sponges.
Many of the most beautiful Alga? are exceedingly minute and
occur in quantity in situations which are not at first obvious, and
which are only found by experience. They occur embedded in a
thin mucus surrounding the stems and leaves of submerged plants,
such as Utricularia, Myriophyllum, Nymphcea, Nnphar, Potcnno-
yeton, Scirpus fluitans, Isoetes, etc. Many of the submerged mosses,
such as Sphagnum contort um, S. plumosum, Amblystegium scorpi-
oides, A. falcatum, A. exannidatum, A. glaitcu'in, Fontinalis anti-

pyretica, Jungermannia inflata, Nardia emarginata, etc., are often

richly covered with these minute Alga?.

Numerous minute Alga? occur along with an abundance of the
Peridiniea? and Copepoda scattered through the surface waters of
lakes, rivers, and other large bodies of fresh water, and constitute
a large proportion of the Freshwater Plankton. The animal and
 Introduction 5

vegetable organisms occurring in the plankton form the food of

most of the smaller aquatic animals, and so, indirectly, form the
basis of the food-material of lacustrine and river fishes. Little is
known concerning the food-value of the freshwater plankton, but
statistics of this nature have been
very carefully compiled with
regard to the Baltic Sea. Brandt states that the chemical com-
1

position of the plankton of this sea in autumn and winter is

"
intermediate between that of " rich pasturage and green lupines.
The proportion of fat is greater than in land products used as
fodder, but in spring the great abundance of Diatoms causes such
a great increase in the amount of ash as to preclude direct com-
parison with land plants. Many of the Algse found in the plankton
are more or less characteristic, some of them being largely and
others entirely surface organisms. The majority of them belong
to the Palmellaceae, Protococcacese, Volvocaceas, Desrnidiaceae and
Bacillarieae. Most of the Desmids of the plankton possess very
long spines or processes which terminate in spines, and in those
species which normally possess long spines the latter are of greater
length when the plants occur in the plankton than when found in
other situations. Some of the Protococcacese and Diatoms have
also acquired long spines. The assumption of this spined con-
dition is to be correlated with their free-floating existence and
their consequent need for greater protection against those animals
of the plankton which feed on Algse 2 .

Algae exist under very varied conditions of temperature. In

temperate and arctic climates many of them can survive prolonged
freezing even when in the ordinary vegetative condition. It is

quite possible to melt out from the ice numerous healthy Algae
which have suffered in no way from their exposure to such a low
temperature. In the arctic and antarctic regions, in the Alps and
in the Andes, there is a snow-flora, consisting principally of Algae
which pass their entire existence on the snow and ice. This
collection of Alga^, which is known as the Cryoplankton,' consists
'

of a few forms which are more or less universal in such situations 3 .

1
Brandt in Wissensch. Meeresuntersuchungen, N. Folge, Bd iii, Heft 2, 1898 ;

consult also American Naturalist, xxxii, Dec. IH'.IS.

2
West & G. S. West, 'Scott. Freshw. Plankton,' Journ. Linn. Soc. Bot. xxxv,
Nov. 1903, p. 554.
3
The most interesting of these Algse are Spfuerella nivalis Sommerfeldt (the Red
Snow plant) and a Desniid, Aneylonema Nordenakioldii, tirst discovered byBerggren
in the snows of Greenland, and afterwards by Lagerheim in the Andes and by Chodat
on Mont Blanc.
6 Introduction

Algae also occur in warm streams, and the vegetation of hot-springs

consists exclusively of
Algae. They can exist in hot water and hot
vapour up to a temperature of 94'5 C. (200 F.) 1 It is worthy of
.

note that the Algae which occur at

very high altitudes, and which
therefore exist at relatively low
temperatures, and those which
inhabit the hottest springs, are, with few
exceptions, species of
Myxophyceae and Bacillarieae.
Some Algae become encrusted with carbonate of lime or with
silica, and play no small part in the formation of the
deposits
which are generally found in the neighbourhood of
hot-springs.
The comparative richness of any district in freshwater
Alga?
depends very largely on its physical geography and on the geo-
logical formations. Mountainous tracts are more prolific than flat
districts, even though many of the larger Algae are absent from
them. Most of the larger filamentous Algae and an abundance of
the commoner unicellular forms are found in
low-lying quiet
waters, but in mountainous areas the filamentous forms are chiefly
representatives of the Myxophyceae and Conjugatae, the presence
of numerous species of the genus
Mougeotia being a marked feature
of such districts, and the unicellular forms are
greatly increased
by the addition of numerous Desmids. If the mountains consist
of the Older Palaeozoic rocks, of Pre-Cambrian rocks, or of rocks of

Igneous origin, there is a surprising numerical increase, not merely

of species but also of individuals and in;
comparison, a mountainous
district of carboniferous limestone or other formation is
distinctly
poor. Thus, the English Lake District, Wales, and certain parts
yield a much greater variety of Algae than
of Scotland and Ireland

any other parts of the British Isles. The poorest area of all is the
fen district in the east of
England.
The most
prolific localities in the British Islands, and perhaps
in the whole of Europe, for freshwater Algae are the small tarns
and peat-bogs which lie in the hollows of the Lewisian
gneiss of
north-west Scotland. The plankton of the larger lakes of this
area is also much richer in the Desmidiaceae than
any which has
been described from elsewhere.
Most of the unicellular Algae and some of the filamentous ones,
unless specially protected as in
many Desmids, are readily taken
1
W. H. Brewer in Amer. Journ. Science, ser. 2, x)i. These Algas were
unicells,
filamentous Algre having been observed
up to a temperature of 85 C (185 F)-
G. S. West in Journ. Bot. 1902, 241.
p.
 Introduction 7

as food by Amoebse, Turbellarians, Oligochaetes, Tardigrades and

Crustacea. The tadpoles of the common frog feed almost exclu-
sively on the larger filamentous Algae, and Bles has recently shown
that the larvae of that most remarkable African frog, Xenopus
Icevis, feed exclusively on the lower Algae
1
A considerable pro-
.

portion of the food of freshwater Lamellibranchs also consists of

livingand decaying Algae.
For the collection of freshwater Algaa a plentiful supply of
small wide-mouthed tubes or bottles will be found most useful.
Small quantities of the larger, strictly aquatic Alga? should be put
into these tubes, care being taken not to overstock the tube. A
small tuft of the Alga with plenty of water is best, and the tube
should not be filled more than three-quarters full. The reason for
this is obvious, as rapid decomposition accompanies overcrowding,
and it frequently happens that collections cannot be examined for
many hours after they have been made, or under certain circum-
stances even for a day or two.
/

To collect the minute Algae that occur attached to larger water

plants, the latter should be removed from the water with
as little
mud as possible, the superfluous water allowed to drain away for a
few minutes, and they should then be gently squeezed over a wide-
mouthed bottle, the issuing water being collected in the bottle.
In the sediment which settles to the bottom of the water in the
bottle will be found numerous unicellular and other small Algae.
Sometimes a hundred or even two hundred species may be obtained
from a small quantity of such material. The Algae of the plankton
are collected from the surface layers of water by means of silken
tow-nets.
If it is desired to keep the plants living for some time they
should be placed in wide-mouthed jars with an abundance of
water, and not exposed to too strong light. Sterile species of the
will often become under
Zygnemaceae and (Edogoniaceae
fertile

these conditions.

1
There can be little doubt that a portion of the food of Ceratodus, the Australian
mud-fish, consists of Algae. An examination of the intestine of this fish (for which
I must express my indebtedness to Prof. Howes), revealed masses of sticks, twi^s,
leaves, fragments of Hepatics, etc., all of which
would have been taken in by the
fish from the muddy bottom of the water in which it lived. This material would
be mostly in a dead condition before being swallowed and it seems to suffer little
change in its passage through the gut ;
but a microscopical examination shows
amongst it the decomposed remains of many kinds of Algae, including thousands
of the empty valves of Diatoms.
8 Introduction

Algae should always be examined in the living state whenever

possible, as some of them are more easily determined when alive.
Many of the Desmidiaceae and Bacillariese, however, especially those
with characteristic surface markings, can only be determined with
precision from the dead empty cells or semicells.
In preserving Alga? for future examination several fluids may
be used. In studying the structure of the cell-contents a 2 4 / o
formalin solution is best for subsequent staining, etc. This is
almost equalled by a dilute solution of picric acid. But if only
the cell-outlines and the structure of the cell-wall are required
then there is no better preservative than a 4 / solution of potas-
sium acetate (containing a trace of copper acetate). An equal
volume of this solution added to the water in which the Alga3 are
living is quite sufficient. A very instructive paper has been
published by Pfeiffer R. v. Wellheim on the methods of prepara-
1

tion, staining, etc., of algse, in which chromacetic acid is largely

recommended for preserving them. Certain reagents such as a
solution of iodine, methylene blue, hsematoxylin, ammonia-car-
mine, etc., are almost essential to all students of Algse. 2 / A
cocaine solution is also very useful for the observation of the cilia
of motile forms.
Some
Algae can be preserved by drying, either on paper or on
slips of mica, but they are always better for purposes of future
examination when preserved in a fluid medium than when dried.
The only Algse that preserve
well by drying are the Myxophycese.
These Algse on soaking out again in water almost regain their
original freshness, and, moreover, retain their original bright
colours.

Algse are best mounted in the fluid in which they have been
preserved, and the best varnish with which to seal them up is
gold-size. Everyone who has had any experience of fluid mounts,
however, knows quite well that if his specimens keep for a long
time it is due more to good fortune than to any other cause.
Many fluid mounts, even the best ones, frequently begin to dry up
by the formation of air-bubbles in the centre of the slide, which
gradually extend towards the periphery. The only explanation of
this is the porous nature of the thin
coverslip.
In examining Algae, in following out their life-history, and in
1
F. Pfeiffer R. v. Wellheim, '

Preparation der Siisswasseralgen,' iu Pringsheirn's

Jahrbuch. t'iir wissensch. Botan. Bd xxvi, Heft 4, Berlin, 1894.
 Introduction 9

identifying them, they should always be carefully drawn

to scale
with the help of a Camera Lucida. This is the surest way of
obtaining accurate measurements of the plants, and also much the
best way of impressing on the memory their diagnostic features.
Great attention should be paid by all students of Algse to their
cytological characters, the structure of the cell-wall, the disposi-
tion of the protoplasmic cell-contents, the form and structure of
the chromatophores, the presence or absence of pyrenoids, etc. It
is the absence of any definite information on these points that

renders many books on Algae almost useless. No student can

acquire erroneous ideas of such common genera as Ulothrix, Tri-
bonema (Conferva), Microspora, and others, if he has once realized
their fundamental cytological characters.

CULTIVATION OF ALG^E.

It is often desirable to cultivate Algse in order to investigate

their life-history and polymorphism. Under cultivation an Alga
is often under abnormal conditions, and as a result, it sometimes

develops strange forms which are quite unknown in the natural

state of the plant. Careful observation of these cultures frequently
affords good evidence towards the elucidation of the phylogenetic

relationships of Algse. Cultures can be made at various tempera-

tures, in water, in sugar solutions of various strengths, or if neces-
sary under damp conditions only. Cultures are most frequently
made in solutions of a nutritive character, but sometimes good
results are obtained in pure water or in weak saline solutions. A
medium containing gelose is very favourable for making pure
cultures of the lower Algse
1
. Klebs and others have emphasized
the usefulness of cultures on gelatine, agar-agar, and other solid
media, and cultivation experiments on damp porcelain plates are
frequently a success. The temperature necessary to obtain the
best cultures varies with different types of Algae, but 20 C. is a

good average temperature.

1
Chodat and Grintzesco in Arch. Sci. Phys. et Nat. x, 1900, p. 386.
 ALG^E.

ALG.E are Thallophytes of a simple or complex structure, and

are of a green, yellow-green, blue-green, red or brown colour.
Most of them live entirely submerged in water and the major
portion of them inhabit the sea. They are found floating freely at
the surface, attached to stones, or as in a large number of the fresh-
water forms, adhering in gelatinous masses to the submerged
portions of more highly organised aquatic plants. A few prefer
damp situations in which they do not become immersed at all, or

only periodically become covered with water.

They are mainly distinguished from the Fungi by the presence
of chlorophyll and consequently by their mode of life. Even in
the red, brown, and blue-green Algse chlorophyll is present, but
the green colour is masked by the presence of other colouring-
matters. As the colouring-matter is usually the same throughout
large groups of these plants which agree in other characters,
particularly in the method of reproduction, they are classified as
follows :

Class 1. Rhodophycece (or the Red Algas), containing a

reddish colouring-matter known as phycoerythrin.

Mostly marine.
Class 2. Phceophycece (or theBrown Algae), containing a
brown colouring-matter known as phycophsein.
Mostly marine.
Class 3. Chlorophycece (or the Green Algae), containing only
the green colouring-matter known as chloro-
phyll. Very largely freshwater plants. The
stored product of assimilation is in almost all
cases starch.
 Freshwater Algce 11

Class 4. Heterokontce (or the Yellow-green Algae), contain-

ing a large proportion of a yellow pigment known
as xanthophyll. The stored product of assimila-
tion is 'a fatty substance. Freshwater.
Class 5. Bacillariece (or the Diatoms), containing a brown
colouring-matter diatomin, which much resembles
the phycophnsin of the brown Algae. Universal
both in fresh and salt water.
Class 6. Myxophyceie (or the Blue-green Algae), containing
a blue colouring-matter known as phycocyanin.
The stored product of assimilation is most probably
glycogen. Mostly freshwater.

By far the greater part of the vegetation of the sea consists of

marine Algae, and with few exceptions these marine forms are of
quite a different nature from the freshwater ones. It is only with
freshwater Algae that this volume is concerned.
Certain Algae are known in a fossil state. These are mostly
Diatoms, the siliceous valves of which are eminently suited for
fossilization, and a few others in which the thallus was encrusted
with carbonate of lime. The majority of other Alga? are of much
too fragile and delicate a nature to become fossilized, and most of
the records of such fossil Algae are of very doubtful value.
Freshwater Algae exhibit a variety of types of structure.
Some of them are unicellular, each plant consisting of a single

protoplasmic unit or energid (i.e. a mass of protoplasm containing

a single nucleus) surrounded and enclosed by a definite cell-wall
(e.g. Desmidiacese, Bacillarieae,
and many Protococcoideae) others ;

are unseptate or ccenocytic plants composed of an aggregate of

protoplasmic units enclosed within a common

cell-wall (e.g. Sipho-

nese, Hydrodictyon); others are incompletely septate plants, each

segment containing a number of protoplasmic units within a cell-
wall, the septation of the plant going on independently of the
divisions of the nuclei (e.g. Cladophorales) ;
others are multi-
cellular or completely septate plants, eachsegment containing one
protoplasmic unit (e.g. Zygnemaceae, Chaetophorales, etc.).

Many of the unicellular forms are solitary cells, but others

occur as colonies, in which the individual cells are more or less
loosely held together in a common mucilaginous envelope,
which
is either secreted by the protoplasm of the cells or is derived from
12 Algce
the cell-walls.This mucilaginous sheath is present in most of the
unicellularand filamentous freshwater Algse, and sometimes attains
a huge development. Its nature was well investigated by Haupt-
fleisch and more recently by Schroder 2
1
.

The
multicellular forms consist of closely connected cells form-
ing a thallus, which exhibits a great variety of form. It may be
spherical (e.g. Coslastrum), filamentous (e.g. Spirogyra, Ulothrix,
etc.), or a flattened expansion (e.g. some species of ColeocJuete,
Protoderma). Sometimes the thallus is differentiated into a
" " " "
root and a thalloid shoot
(e.g. Botrydium, Rhizoclonium,
3 " "
(Edogonium, Spirogyra, etc. ), but the root is in all cases merely
an organ of attachment and is more correctly called a hapteron.
The cell-wall always consists largely of. cellulose, and is some-
times delicate, sometimes of considerable thickness and strength,
being cuticularized or even silicified, but it is rarely, if ever,
lignified. It often becomes gelatinous in its outer layers.
The
principal colouring-matter of the cell is usually arranged
in definite parts of the protoplasm known as chromatophores. A
single cell may contain one or many chromatophores. If the

chromatophores contain the green colouring-matter chlorophyll,

they are known as chloroplastids (or chloroplasts} if they contain ;

some other colouring-matter they are termed chromoplastids (or

chromoplasts). Plastids are present in all groups of Algoe, but
those present in the Myxophycese are of a very primitive character.

Chromatophores, particularly chloroplasts, often contain pyrenoids,

which consist of colourless masses of proteid substance. The
central mass of the pyrenoid is a proteid substance of crystalloidal
character which bears a great resemblance to an aleuron grain.
It is sometimes angular, sometimes rounded, or it may be quite

irregular in its outward form, and is often surrounded by an

amylaceous envelope or coat of starch. The latter sometimes
becomes lobed and penetrates into the chromatophore to such an
extent that its existence appears doubtful. In Spirogyra the
envelope of amylaceous material round each central mass (or
pyrenocrystal) is in the form of a number of grains of starch. On

Hauptfleisch, Zellmenibran und Hiillgallerte der Desmidiaceen,' Mitteil. aus

1 '

d. Naturwiss. Vereine f. Neuvorpommern und Rugen, 1888.

- '
Schroder, Uutersuchungen iiber Gallertbildungen der Algen,' Verhand. des
Naturhist.-Med. Vereins zu Heidelberg, Bd vii, Heft 2, 1902.
3
Borge, Ueber die Rhizoidenbildung,' Upsala nya Tidnings Akteb. Tr. 1894
'

West and G. S. West in Ann. Bot. vol. xii, March 1898.

 Vegetative multiplication 13

the division of a cell the pyrenoids usually divide equally. Some-

times a pyrenoid in a well-nourished cell multiplies by division
without any division of the cell or the cell-nucleus. In a badly
nourished cell, the amylaceous portion first disappears and then
the crystalloidal part. The pyrenoid is thus a store of reserve
food-material, and arise quite spontaneously without the
it
may
previous existence of pyrenoids in the cell.
Almost all Algae are holophytes; that is to say, they are them-
selves able to elaborate organic material from the mineral and
other inorganic substances found in the water, or in some instances,
in the atmosphere, in which they exist. The chlorophyll found in
the chromatophores of the cells arrests certain rays of light, the
of which is utilized by the living for the con-
energy protoplasm
struction of organic substance from the inorganic materials taken up.

Algae absorb a relatively large amount of mineral food sub-

stances, particularly nitrates, from the water in which they live.
It has been assumed that the presence of nitrates in abundance is

necessary for the prolific growth of Alga?, but it is certainly true

that these plants occur in quantity in water which is relatively

poor in nitrates. Whipple and Parker state, as a result of experi-

1

ments on the occurrence of small chlorophyll-bearing organisms in

the waters of lakes, that the presence or absence of carbonic acid is
one of the fundamental factors which influence the growth of Algae.
The tropical Algae of the genera Phyllosiphon and Cephaleuros
are partial parasites, and a few have already been mentioned as

symbiotic with other plants or even with animals.

The growth of the thallus may be apical or intercalary. In
many Algae by the repeated division of a single apical cell, or
it is

by a series of marginal cells, as in the expanded thallus of Coleo-

chwte. In many of the filamentous Algae with intercalary growth
all the cells of the thallus are meristematic and undergo division
(e.g.Zygnemaceae, Ulotrichacese).
Vegetative multiplication occurs in the unicellular forms by
ordinary cell-division or fission, and in many of these plants it is
the only method of increase. The division may take place in one
direction only (e.g. Aphanothece, Gloeothece, Stick ococcus), in two
directions in one plane (e.g. Tetraspora, Merismopedia), or in all
directions of space (e.g. Gloeocystis, Glceocapsa, and many others).

1
Whipple and Parker, in Trans. Amer. Micr. Soc. May 1902.
 Algw
In the Desmidiacese, in which the cells generally exhibit a more
or less deep median is only in one direction,
constriction, division
and it is
brought about by the development of two new half-cells
(or semicells) between the old halves. So that each of the indi-
viduals formed after one division consists of an old and a new
half-cell. In many unicells the division is accomplished by the
formation of daughter-cells within the mother-cell. The daughter-
cells arerejuvenized and metamorphosed portions of the original
mother-cell and are enclosed in the old wall of the mother-cell.
Sometimes the daughter-cells are set free by the bursting of the
wall of the mother-cell, but it often happens that the old wall of
the mother-cell swells up and remains as an outer, wider coat to
the daughter-cells. Successive generations of cells are produced
in the same manner inside the enlarged walls of the mother-cells
until quite a colony is formed inside the swollen cell-wall of the

original mother-cell (e.g. many Protococcoideae and Chroococcaceas).

In many types of lower Algae there is no definite line to be drawn
between this so-called free cell-formation and the ordinary vege-
1
tative division . In certain of the Protococcaceae two or four
daughter-cells arise in a mother-cell, and at the time of their
escape from the parent-cell they possess the exact form and external
peculiarities of the parent these have been termed autospores.
;

Increase of the filamentous Alga? by new

cells occurs in
divisions, the septa
being always transverse to the length. In the
(Edogoniaceae the method of cell-division is somewhat specialized
and a description of it is given under the family.
On injury to the filamentous coenocytic Algae septa usually
appear cutting off the injured part. The injured portion soon dies
away, and if it happens to be in the median part of a filament,
two filaments are thus set free. This occurs frequently in
Vaucheria, and if the injuries to one filament are numerous, all
the injured parts are sometimes cut off, the intermediate uninjured
portions developing into new filaments on being set free.
The reproduction of freshwater Algae is brought about in a
great variety of ways, most of the plants exhibiting both an
asexual and a sexual mode of reproduction.
Asexual reproduction. In some cases special non-motile
cells develop into cysts or unicellular gemmae (e.g. Zygnemaceae

1
Cuoclat in Ann. Bot. 1897, p. 102.
 Asexual reproduction 15

they are also specially cutoff in the Vaucheriacese), and in the

filamentous Myxophycese hormogones or multicellular gemmae are

formed. Sometimes single non-motile cells are produced, which
have been termed by Wille akinetes when they are formed without
rejuvenescence and aplanospores when formed by rejuvenescence
(e.g.Chsetophorales, GWfervales, Conjugatse). Many of these non-
motile asexual spores, which may be either akinetes or aplanospores,
rest for considerable periods before germination and are known as

hypnospores (or sometimes as hypnocysts). Asexual reproductive

organs are known
as gonidangia when borne on the gametophyte

generation and sporangia when borne on the sporophyte. A

a
sporangium (or gonidangiurn) as a rule gives origin to a number
of spores (or gonidia), but in Vaucheria, (Edogonium, and some of
the Ulotrichacese only one gonidium is formed in the gonid-
angium.
Very often the gonidia consist of motile cells which receive the
name of zoogonidia (or zoospores). One of the most frequent
methods of asexual reproduction is by means of zoogonidia, which
are found in all groups of freshwater
Algas except the Conjugates
(the largest order of the Chlorophyceas), the Myxophycese, and the
Bacillarieae (or Diatoms). Zoogonidia are small masses of proto-
plasm formed singly by the rejuvenescence of the entire contents
of a or more frequently in numbers by free cell-formation.
cell,

They are not possessed of a cellulose wall, but are furnished with
one (?), two, four, or many cilia, with one or more chromatophores,
and often with one or two contractile vacuoles. The cilia are
usually disposed towards one end or one side of the zoogonidium
and their rapid vibratile movements cause it to swim quickly
through the water. A red or brown pigment-spot is very often
present. After a time the zoogonidium comes to rest, the cilia
disappear, the protoplasm secretes a cellulose wall, and the zoo-
gonidium develops into a new plant.
Sexual reproduction. Reproduction by the union of male
and female elements, or gametes, is fairly general throughout the
Algae, but it is entirely absent in the Myxophyceae, in some of the
unicellular Protococcoideaa, and in the Syngeneticse. Sometimes
the gametes are clearly differentiated into male and female ele-
ments, but in other cases sexual differences are scarcely appreciable.
The following is a summary of the sexual methods of reproduction
met with in the freshwater Alga^ :
16

I. Sexual reproduction by isogamous gametes (isogametes), or

precisely similar sexual cells which undergo the process of con-
jugation.
a. Gametes ciliated, known as planogametes or zoogametes,
set free, and on conjugation forming a zygospore (or
zygote} (e.g. Ulotrichacea?, Ulvacese, Trentepohliaceae,
Pandorina).
b. Gametes not ciliated, known as aplanogametes.
i.
Conjugation forming a zygospore which after a period
of rest develops directly into a new gametophyte

(e.g. Bacillariea? and Desmidiaceae, in which the

gametes are set free ; Zygnemeae, in which the
gametes are not set free).

ii.
Conjugation forming a zygospore which immediately
develops a rudimentary sporophyte with one spore
(e.g. Mesocarpeae).
II. Sexual reproduction by heterogamous gametes (hetero-
gametes), or clearly differentiated sexual cells which undergo the
process of fertilization.
a. Oogamous heterogamy The female organ is an oogonium
:

containing an oosphere. The male organ is an antheri-

dium in which are developed motile, ciliated antherozoids
(or spermatozoids). The result of fertilization is the

production of an oospore (e.g. Yaucheriaceae, (Edogo-

niales, certain of the Chaetophorales and Cladophorales,
and some of the Volvocacese).
b.
Carpogamous heterogamy: The female organ is aprocarp
(consisting of carpogonium and trichogyne) with no
specially differentiated female cell. The male cells are
non-ciliated spermatia (or pollinoids}. Fertilization
results in the development of a cystocarp (or sporocarp)
with contained spores known as carpospores (e.g.
Rhode >phyces?.).

The sexual organs of those Algas with similar sexual cells are
termed gametangia. If the Alga? are unicellular then the cell
itself becomes the gametangium (e.g. Desmidiaceae, Bacillariese),
and in the multicellular and ccenocytic forms the ordinary vegeta-
tive cells become the gametangia (e.g. Zygnemaceae, Chastophorales,
 Sexual organs 17

Hydrodictyori). In the whole of the Conjugatae the gametangium

gives origin to only one aplanogamete, but in other Algae it is
more usual for several gametes to arise from one gametangium.
Planogametes, such as are found in the Ulotrichaceae, are pear-
shaped bodies with the chromatophores more or less confined
towards the broader end, the narrower end being colourless. Two
cilia are inserted at or near the narrow, colourless end, and a red

pigment-spot is frequently present. They exhibit active move-

ments for a longer or shorter period of time and finally conjugate,
each pair coming into contact by their colourless poles.
In those Alga? with dissimilar sexual cells the female organ
consists either of a single cell or a coenocyte known as the oogonium,
which is usually more or less spherical (e.g. CEdogoniacea?), some-
times attenuated into a beak (e.g. Vaucheria), or produced at the
apex into a long, narrow tube, the trichogyne (e.g. Coleochaetaceae).
An oogonium usually contains a single female cell or oosphere
(e.g. Vaucheriacere, GEdogoniaceae), but sometimes there are many
oospheres present (e.g. Sphseropleacea3). An oosphere is generally
a spherical cell containing chromatophores, and often with a clear,
colourless area at one side known as the receptive spot. It is at this

spot that the antherozoid enters during the process

of fertilization.
In the Rhodophycea? the female organ is usually a inulticellular
structure (in the freshwater Red Alga? it is unicellular) termed a

procarp, which is divisible into two portions, a carpogonium and

a trichogyne.
The male organ is known as the antheridium. It is usually

unicellular, but in (Edogonium may it consist of one or of many

cells. Each antheridial cell often gives rise to quite a number of
male cells (spermatozoids or antherozoids), but in the (Edogoniacese
it gives rise to two, and in the ColeochaetaceaB and most of the

Rhodophyceas to only one. Antherozoids frequently resemble the

asexual zoogonidia, but are usually smaller. They are commonly
pear-shaped, but may be elongate and almost rod-like. They
possess two cilia which are generally inserted at the pointed end,
but laterally in Volvox, Vaucheria, etc., and in (Edogonium theiv
is
quite a tuft of cilia at the narrower end. They are as a rule
faintly coloured and often possess a red pigment-spot.
Sexual organs have riot been observed in many of the Pro-
tococcoidea? and in the Syngeneticae, and are apparently entirely
absent from the whole class of the Myxophycea?.
w. A. 2
 18 Algce

Many Algae exhibit an alternation of generations in their life-

history. In those forms in which neither asexual nor sexual
reproduction is known this alternation of generations is, of course,
absent. Other Algae exhibit slight indications of an alternation
of generations. Thus, in a large number of the Chlorophycese, the
sporophyte generation is represented by the zygospore. This
zygospore can be described as a unicellular sporangium which often

gives rise to two or four zoospores. Each zoospore, on coming to

rest,germinates and produces the gametophyte generation. In
the Mesocarpeae the isogamous gametes conjugate in a connecting-
tube between the gametangia, and the zygospore immediately
produces a rudimentary sporocarp consisting of a variable number
of cells, one cell of which is the carpospore. This is the sporophyte
generation. The carpospore, after a period of rest, germinates and
gives rise to the new gametophyte generation. In the Rhodo-
phyceae there is a well-marked alternation of generations.
In the Algge the gametophyte is the important generation ;

'

in fact, the 'plant is the gametophyte ; but as one proceeds higher

in the scale of vegetable life there is a great reduction of the

gametophyte accompanied by a corresponding increase in the

development of the sporophyte, until in the Phanerogams the sporo-
phyte becomes the 'plant' and the gametophyte is parasitic on it.

POLYMORPHISM.

Polymorphism occurs in most families of Algae. All those Algae

which exhibit an alternation of generations are polymorphic, and
some species appear to possess several different vegetative forms.
It is very doubtful, however, if polymorphism occurs in Algae to
the extraordinary extent advocated by some authors. Sirodot has

proved the occurrence of several types of structure in the genera

Batrackospermum and Lemunea, and it is fairly evident that the
plants at one time described under the genera Prasiola, Schizo-
(jonium and Hormidium are different vegetative forms of one
genus. Polymorphism is well illustrated in Botrydiutu, and
numerous striking instances could be mentioned of other Alga? in
which it occurs, notably in the common genus Pleurococcus.
Hansgirg, and following in his footsteps Wolle, have endeavoured
to prove (on insufficient evidence) the existence of extraordinary

polymorphism in Algae, relegating most of the unicellular Algae as

 Polymorphism 19

mere stages in the growth of higher forms. It is necessary, how-

ever, to remark that a great many loose statements have been made
on this subject, statements which are supported by no direct or
conclusive evidence. Most bf the remarks have been based upon the
fact of the occurrence together, in one matrix, of various stages of
different plants, and to the assumed identity of certain normally
unicellular plants with unicellular stages in the life-history of

higher organisms. Undoubtedly in the case of the Myxophyceae

many different forms are met with in one gelatinous matrix, and
these are of the most confusing nature. Many of the higher blue-
green Alga? do certainly pass through stages which resemble very
much some of the so-called unicellular species, but there is equally
no doubt that careful observation frequently proves that this is a
resemblance and not an identity. In some instances it may be quite
true that a blue-green form which has been accepted as a species
in the past is really a stage in the development of another form, but
that does not necessarily prove that every blue-green Alga exhibits
wide polymorphism and that every form met with is only one stage
in some complex life-history. Similarly, in the Chlorophyceae,
is frequent, but because species of Chcetophora or
polymorphism
Myxonema (Siigeoclonium) at one period of their existence regularly
'

and normally break up into

'

Palmel la-like forms, it does not

follow that every aggregate of unicells such as Glceocystis, Palmella,
etc., is merely a stage in the development of Ghcetophora, Myxonema,
or some other allied plant. The observation of the polymorphism
of higher and lower types of Algae, both in nature and under
cultivation, is, however, the surest and best way to discover their
affinities, and in many cases furnishes direct evidence as to the
phylogeny of the plants in question. Thus, the fact that Myxo-
nema assumes under certain conditions a Palmella-siate,' much
'

in no way specifically
resembling species of the genus Palmella but
related to them, is one of the primary reasons for regarding

Myxonema (and therefore the Chastophoracese) as having been

derived from the Palmellace*.
Without question many of the Alga3 referred to the order
Protococcoidese have a much more direct relationship with fila-
mentous green Algae, particularly with the Chsetophorales, than is
indicated by their present systematic position. As an instance,
the genus Stichococcus, described as a unicell, un-
by Nageli
the Ulotrichacea? and is connected by
doubtedly belongs to family
22
20 Alga'.

many intermediate stages with species of Kiitzing's genus Glceotila,

and even with species of the genus Ulothrix. This was first
2
definitely shown by Gay and afterwards emphasized by Klercker
1
.

But, although in some instances this is the case, and various

'

genera of the group Protococcoideae have to be transferred to

'

other groups to which they more rightly belong, many forms still
'

have to remain in the old group Protococcoidese until more is

'

known concerning their life-histories and affinities. For instance,

it has been asserted that species of the genus Tetraedron Kiitz.
1845 (= Polyedrium Nag. 1849) are merely stages in the develop-
ment of Pediastrum, but this is now known to be erroneous 3 ,

and that even if certain forms are developed in the life-cycle of

Pediastrum which much resemble species of Tetraedron, yet the
latter genus is quite distinct and has a life-history of its own. It

is, however, most probable that Pediastrum has been evolved

from
Algae of the nature of Tetraedron, and one of the connecting links
has been discovered in a genus recently described by Lagerheim
under the name of Euastropsis.
As another example of erroneous conclusions arrived at from
insufficient evidence, there is the case of the genus Chantransia.
'

The discovery by Sirodot 4 of the protonema-stage of Batracho- '

spermum was regarded by many people as a sufficient proof that

species of Chantransia were merely asexual forms of Batracho-
spermum. This was entirely due to the mistake of confounding
the sporophytic shoots of Batrachospermum (and Lemanea) with
certain true species of Chantransia which they much resembled.
5
Murray in commenting upon this, says that to speak of the
,

"
" "
Chantransia-formsof these genera means no more than if the
'

protonema of a moss were to be called its Conferva-farm or

'

the prothallus of a fern its Liverwort- form.' These growths of

'

Lemanea and Batrachospermum have nothing to do with the valid

generic type Chantransia"
In conclusion, it seems well established that the higher types
of Algae have originated by gradual evolution from the more lowly

1
Gay,
'
Recherches sur le developpernent et la classification de qnelques Algues
Vertes,' Paris, 1891.
2
Klercker,
'
Ueber zwei Wasserformen von Stichococcus,' Flora, 1896.
3
Tromso Museums Aarshefter, 17. 1894.
Cfr Lagerheim in
4
Sirodot,
'
Sur
le developpernent des Algues d'eau douce du genre Batracho-
'

spermum, Paris, 1875.

5
G. Murray, An Introduction to the Study of Seaweeds,' London, 1895, p. 208.
'
 Phylogeny 21

types, but the fact must not be overlooked that these lowly types,
although they may have undergone many modifications, still

persist,and great care should be taken not to confound them with

those stages in the life-histories of the higher types which present
so many resemblances to them.

THE PHYLOGENY AND CLASSIFICATION OF THE

FRESHWATER ALG.E.
The researches and discoveries of the last few years have
certainly thrown much light on the affinities of many genera and
families of Alga and constitute a very great advance in our
1
,

knowledge of the phylogenetic relationships of these plants. It is

by no means an easy task to give even a mere outline of the
suggestions which have at different times been put forward as to
the evolution of freshwater Alga?, but one derives great assistance
from two recently published papers, one by Chodat and the other
1

by Blackman containing not only a summary of much of the. work

2
,

bearing on this difficult problem of phylogeny, but putting forward

some well-founded suggestions as to the same.
In the succeeding brief account of the evolution of freshwater
Alga? I have followed very largely the suggestions of Borzi,
Blackman, Bohlin and others, with certain alterations based upon
my own
3
experience .

Taking first the Chlorophycea? or green Alga?, which a few

years ago were in a chaotic condition, we find that this chaos
has
been greatly reduced to order and that the affinities of many of
these plants have been clearly demonstrated. The four groups
of the Confervoidese, Conjugate, Siphonea? and Protococcoideae, into
which the green Alga? have been usually classified, must be con-
siderably modified in view of recent researches.
The Conjugate
and the Siphonea? will remain as distinct and natural orders of the

green Alga?, the former chiefly by reason of their reproduction

and
the latter on account of their ccenocytic structure, but the Con-
fervoidea? and ProtococcoideaB were unquestionably unnatural

Chodat, 'On the Polymorphism of Green Algae and the Principles of their
1

Evolution,' Ann. Bot. xi, 1897.

2
F. F. Blackman, 'The Primitive Algre and the Flagellata. An Account of
Modern Work bearing on the Evolution of the Algas,' Ann. Bot. xiv, I'.tno.
3 In
'Lectures on the Evolution of Plants' by D. H. Campbell (Macmillan
Company, New York, 1889), there is a chapter on Algae (pp. 4879) with a scheme
of evolution (p. 79), but the latter appears to be largely based upon erroneous
conceptions of the relationships of these plants.
22 Alg<c

groups which could no longer be tolerated in the sense in which

they were originally proposed.
Chodat, from observations on the lower green Alga?, both in
a state of nature and in cultures, traces the principal groups
of the Chlorophycese back to the Palmellaceas, one of the lowly
families the order Protococcoidese.
of He recognizes three
important tendencies which rule the lower green Algse: (1) the
zoospore-condition, the other two conditions being only transient ;

(2) the sporangium-condition or unicellular motionless stage, the

other conditions being realized accidentally; (3) the Tetraspora-
stage, where non-motile cells are connected by regular cell-walls
at right angles.
Blackman follows somewhat on these lines, but he considers,
along with others, that all the tendencies of the lower Algce have
had an origin in the motile unicellular Chlamydomonads. Among
the families of lower Algae constituting the group of the Proto-
coccoidese,he observes three divergent vegetative tendencies :

(1) a Volvocine towards

the aggregation of motile
tendency
vegetative cells into gradually larger and more specialized motile
true co3nobia (2) a Tetrasporine tendency towards the formation
;

of aggregations by the juxtaposition of the products of septate

vegetative cell-division to form non-motile organisms of increasing
definiteness and solidarity; (3) an Endosphaerine tendency towards
the reduction of the vegetative division and septate cell-formation
to a minimum. The simplest forms which exhibit any one of these
three tendencies seem clearly to diverge from species of the genus
Chlamydomonas, and these motile organisms must be regarded
as the real primitive form of green plant and the foundation
stone, so to speak, of the vegetable kingdom. Of late much work
has been done at the genus Cltlami/domonas by Goroschankin,
Franct^, Klebs, and Wille, and now the genus is brought
Dill,
into more prominence. It has been found to contain some
still

twenty-nine species which are remarkable for the constancy

of their cytological characters. Unfavourable conditions produce
'
in this genus the Palniella-cofidition.' This is the beginning of
a vegetative non-motile existence such as predominates in the
Palmellacese. In the latter family the cells at intervals in their
life-history escape from their walls, develop cilia, and return to the
motile state as zoogonidia. Blackman remarks that the "formation
of zoospores is then nothing but reversion to an ancestral type of
 Phylogeny 23

vegetative existence for a biological advantage, and all the vegetative

phylogenetically a new intercalation
existence of the higher Alga' is
into the life-history of the motile
Chlamydomonad which is perma-
nently in the zoospore condition, though walled, and in which
zoospore-formation and vegetative cell-division are one and
indistinguishably the same thing."
Chlamydomonas itself has had an origin from the Proto-
mastigina one of the five subdivisions of the Flagellata proposed
by Klebs and two instances of the connecting forms are found
in the organisms known as Polyblepharis Dang, and
Chlorogonium
Ehrenb. Many of these lowly Flagellates are green, but others
are colourless saprophytic organisms, and in some either a
saprophytic or holophytic nutrition may be carried on, depending
on whether the organism is well fed or not 1 In the same way .

that green organisms occur among the Flagellates so do colourless

forms occur among the lower Algae. One species of Chlamydomonas
Chi. hyalina is
always colourless and saprophytic, and Dangeard's
researches into Polytoma have likewise shown that the small group
of organisms of which Polytoma uvella Ehrenb. is the best known

representative should perhaps be regarded as a saprophytic

sub-family of the Volvocacese which have probably evolved from
a green organism of the Chlamydomonad-type. The various
Flagellates such as Eugl&na and others belonging to the Eugle-
noidina, have given origin to no organisms of preponderating
plant-characters.
The Volvocine tendency in the Chlamydomonad-type has
caused the evolution of a series of
organisms gradually of

increasing complexity, which constitute the Volvocacea?. These

are genera which practically consist of coenobia of cells of the

Chlamydomonad-type. The genus Goniam is perhaps very little

removed from Chlamydomonas except in the possession of a four-
celled or sixteen-celled colony. The highest development reached
isthe Volvox-colony, in which there are highly developed oospheres
and antheridia. Between the isoganious Chlamydomonas or
isogamous Gonium and the highly differentiated heterogamous
condition of Volvox, there are two intermediate stages in Pandorina
and Eadorina.
The Tetrasporine tendency in the Chlamydomonad-type first
resulted in the production of a series of forms in which vegetative
1
Zumstein in Pringsheim's Jahrbtich. fiir. wiss. Botan. xxiv, 1899.
24 Algcc

cell-division gradually replaced the formation of zoogonidia as the

chief method of multiplication. The group of Alga evolved
first

in this direction was the lowly family of the Palmellacea?, in which

directions of space as in
grouped together either in
the cells are all

Palmella, or regularly disposed in one plane as in Tetraspora.

The cells are enveloped in a general mucous envelope formed by
the confluence of the special gelatinous cell-walls, and in every
stage the cells on leaving the envelope are capable of swarming as
zoogonidia. It is to this family of the Palmellacea? that we must
look for the origin of most of the other families of green Algae.
The Endosphasrine tendency in the Chlamydomonad-type has
given certain plants in which vegetative cell-division is
rise to
the
absent, the multiplication of individuals taking place only by
formation of zoogonidia or gametes. These plants belong to the

family Endosphgeraceaa and are almost strictly

unicellular.

So far, then, the phylogeny of the green Alga3 can be illus-

trated by the following simple diagram :

Volvocaceas
A
Endosphaeraceae

Palrnellaceae

Chlamydomonas

The Volvocine tendency has resulted in no higher development

than the Volvox-co\ony, but a very reasonable suggestion has been
made by Blackman as to the origin of the Siphonese by a further
development of the EndosphaBrine tendency and formation of a
thallus, which, although essentially coenocytic. is structurally uni-
cellularand lacks the solidity acquired by septate cell-division.
He remarks that "nothing appears to have been evolved from it
of higher status than an Alga. While the Tetrasporine tendency
has given rise to all the higher green plants, the Endosphaerine
 Phylogeny 25

has only succeeded in producing the elaborate but puny mockery

of them which we find in Gaulerpa"
It is now necessary to trace the further developments from the

Palmellacea?, which family was the first result of the influence of

the Tetrasporine tendency on the Chlamydomonad-type.
The Protococcacese is a group which has been gradually evolved
from the Palmellacese by the direct production of the unicellular
motionless stage with a firm cell-wall as the principal state of the
plant, the zoogonidia and the Tetraspora-skSige being only
tran-
sient conditions in the life-history, and often absent. In the lower
forms of this family the cells are globose with firm cell- walls, and
alltheir reproductive processes show a marked tendency to trans-
form the motile elements into resting spores. In some of the
other forms of the family the cells exhibit great variety of form
(e.g. Oocystis, Nephrocytium, Kirchneriella, Chodatella, etc.), and
reproduction is
largely by a type of spore termed by Chodat an
autospore.Such spores are usually produced in fours inside the
and at the moment of their liberation they possess
mother-cell,
the exact form and external peculiarities of the mother-cell.
Sometimes the autospores are quite free after their liberation
(e.g. Lagerheimia, Oocystis), but at other times they are sur-
rounded by a gelatinous envelope (e.g. Kirchneriella, Nephrocy-
tium\. When the autospores are united together in a colony at
the time of their expulsion an auto-colony is produced. Such is
the usual method of multiplication of Scenedesmus. Forms of the
nature of Ccelastrum and Sorastrum have originated from the
lower Protococcaceffi with autospores, the latter being grouped
together into a globular auto-colony.
The position of the two genera Pediastrum
and Hydrodictyon
is still
very doubtful. I am inclined to agree with Chodat that
the resemblance is due to convergence rather than to a common

origin. In Pediastrum the swarming stage is outside the mother-

cell and the new cosnobium arises by the apposition of motile

zoogonidia. In Hydrodictyon the new coenobium also arises by

the apposition of zoogonidia which have become quiescent, but is
formed inside the mother-cell. Both genera consist of coenobia
of ccenocytes, and for the present they are best kept in the

separate sub-families of the Pediastreae and Hydrodictye;c.

Among the Chretophorales it is clear that the Ulotrk'hare.-r
have had a from the Palmellacere through such forms
direct, origin
26 A Igce
as SticJiococcus, Hormospora, Radio/Hum, Gloeotila and Geminella,
forms in which simple loosely connected series of cells occur em-
bedded in a gelatinous envelope.
The Chaatophoracea? are further specialized forms of the Ulo-
trichacese,which are distinguished mainly by their branched habit.
The polymorphism exhibited by the Chjetophoracese, and the deve-
lopment of zoogonidia and formation of resting-spores, also indicate
a close connection with the Palmellacese, but probably through the
Ulotrichacese. Chodat considers the genus Pleurococcus as one type
of the Chaetophoracese which has been reduced owing to its existence
as a lichen-gonidium, but this is a statement I cannot agree with.
The Ulvales and the Schizogoniales are parallel groups, each
of which has probably had a separate origin from the Proto-
coccoideae. The Ulvacese, especially such forms as Monostroma
and Ulva, have most likely originated from Palmellaceous Alga?
of the nature of Tetraspora.
The genus Microspora is difficult to relegate to its proper
place in a classificatory scheme. It is the sole representative of
the family Microsporaceae and its characters mark it off sharply
from other green Alga?. It may have originated from some aber-
rant form of the Ulotrichacea?, but its affinities are at present
doubtful. It is possible that the forms placed in the genus
Rhizoclonium have originated from Microspora, and by a further
specialization the genus Cladophora
has been produced. The
Cladophoracese (and therefore the Pithophoracese) may thus have
had an origin from the Microsporacea?. Bohlin has recently
1

transferred the Cladophoracese to the order Siphonea? owing to

the coenocytic nature of the segments of the thallus, but the
of opinion. to the
validity of such a change is a question Owing
many points of resemblance between Hydrodictyon and Cladophora,
the former genus may possibly be a degenerate form of the Clado-
phoracese. Since 1897- I have regarded the Cladophoracese as a
distinct family of Chlorophyceae, in close proximity to the Siphoneae
and far removed from the Ulotrichacea?, yet I hardly see the justi-

fication for its inclusion in the Siphonese. I think it better to

place Cladophoracese and Pithophoracese, along with

the the

Sphseropleacese, in a separate order, the Cladophorales.

Bohlin, 'Utkast till de Grona Algernas och Arkegouiaternas Fylogeni,' Akad.

1

Afhandl. Upsala, 1901.

2
W. & G. S. West in Journ. Boy. Micr. Soc. 1897, p. 475.
 Phytogeny 27

Luther and Bohlin have recently advocated considerable

changes in the classification of the Green Algse, most of which
have been rendered necessary by the abolition of the old artificial
'

group of the Confervoidege.' I agree with Bohlin in the estab-

'
lishment of the order Microsporales,' even though it appears to
be giving undue prominence to a small group of more or less
insignificant Algse, because species of the genus Microspora are
referable to no other order of green Algse. Likewise, the CEdo-
goniacese require placing in a separate order because of their
anomalous characters.
Several of the recent students of freshwater Alga? have
attempted to show that all the main groups of the Chlorophycese
have had a separate origin from unicellular, motile, ciliated or
flagellated ancestors. This is no doubt a very helpful idea, but
like many other such ideas it can easily be carried too far. It

appears most probable that certain groups of green Algse have had
a direct origin from ciliated or flagellated unicells, but that in itself
is no proof that other groups have had a similar origin. There is
not a shadow of evidence in support of the direct and individual
origin of the Microsporacese, the Conjugata?, the Vaucheriacese, the
(Edogoniacese or the Cladophorales in fact, there is every reason ;

to suppose that at least of these groups have originated from

some
previously existing filamentous forms.
The origin of the Conjugate seems very uncertain. Black-
man, in his scheme of evolution and Bohlin have both suggested
2 1
,

an origin from the unicellular motile Chlamydomonad-type, and

therefore directly from the Flagellata. To my mind this shows a
lack of experience of the Conjugata? as a whole, and particularly
of the family Desmidiacese. Whatever the true origin of the
Conjugate it cannot have been direct from Flagellate forms.
Presumably the first Conjugates which would arise from such
motile unicells would be themselves unicells, or loose aggregates of
cells. is exactly the condition found in the Desmidiacese
Now, such ;

but has been clearly shown 3 that the Desmidiacese is unques-

it
from
tionably a family of Conjugates derived by retrogression
filamentous ancestors, and therefore, they cannot by any possible
means have had a direct origin from unicellular motile organisms.

1 2 22.
Blackman, I.e. p. 684. Bohlin, I.e. p.
3
W. & G. S. West in Ann. Bot. xii, 1898, p. 55 ;
G. S. West in Journ. Linn.
Soc. Bot. xxxiv, 1899, pp. 40941.").
28

It appearsmore probable that the early Conjugates were filamen-

tous forms and that they originated from some other order of
green Algae, corning to an abrupt conclusion soon afterwards.
Nothing higher was evolved from them, but the group of the
Desmidiacese became sharply marked off from the rest of the
Conjugates owing to great specialization. The specializing ten-
dency was in the direction of a remarkable increase in the com-
plexity of morphological characters, and this was accompanied by
degeneration of sexual differences.
Perhaps there may be an affinity between certain of the Con-
jugates and the genus Microspora, as the resemblance between
such species as Microspora Lofgrenii Nordst. and Zygnema 1

pachydermum West is most striking; the cell- walls are thick

2

and exhibit the same structure, and in both plants precisely

similar aplanospores are formed in identically the same manner.
It is now necessary to consider a group of Alga3 for which
Borzi 3 proposed the name of the Confervales.' In this order he
'

included a number of Algse which had previously been scattered

amongst various groups of the Chlorophycese. The characters of
the group are based upon the structure of the which contains cell,

parietal discoidal chromatophores of a yellowish-green colour and

without pyrenoids. Even the zoogonidia possess discoidal chroma-
tophores of a yellowish-green colour and two unequal cilia (some-
times one ?). The plants may be unicellular, coenocytic or multi-
cellular, and include amongst others the following genera :

Ophiocytium, Characiopsis, Chlorothecium, Mischococcus, Tribonema

(Conferva), Botrydiopsis and Botrydium. Bohlin 4 in 1897 con-
clusively demonstrated, by an exhaustive study of the structure of
the cell-wall, the close affinity which exists between the genera
Ophiocytium and Tribonema (Conferva); and, in addition, in the
earlier stages of development these two genera, one of which is
unicellular and the other multicellular, much resemble each other 5 .

The order Confervales is subdivided into three families: (1) Tri-

bonemacese, which includes Tribonema (Conferva}, Ophiocytium,
1
Nordstedt in Botaniska Notiser, 1882, p. 55 ;
W. & G. S. West in Journ. Bot.
Febr. 1897, p. 34.
-
West in Journ. Linn. Soc. Bot. xxx, 1894, p. 266, t. xiii, f. 1 16.
3
Borzi in Boll, della Soc. ital. dei Microscop. i, 1889.
4
Bohlin in Bihang till K. Sv. Vet.-Akad. Haudl. 1897, Bd xxiii, no. 3.
5
Bohlin I.e. t. ii, 5456: Wille in Ofvers.
47, 51, 52,
f. af K- Vet.-Akad. Forh.
1881, no. 8, t. ix, f. 15, 17, 18,2126; G. S. West in Journ. Bot. Mar. 1899, p. 106,
t. 394, f. 1822.
 Phytogeny 29

Bumilleria, Botrydiopsis, Chlorobotrys (2) Chlorotheciacece, which

;

includes Mischococcus, Peroniella, Stipitococcus, Characiopsis,

Ghlorothecium (3) Botrydiacese, including Botrydiiun.
; Now,
amongst the Flagellate organisms there exists a genus described as
Vacuolaria by Cienkowski which possesses yellow -green discoidal
1
,

chromatophores without pyrenoids and this organism, as in the

;

case of theChlamydomonad-type, may very possibly be the start-

ing point of the Confervales. Lagerheim discovered another
organism in 1897 which was further worked out by Bohlin- and
named Chloramceba. It is of a similar type to Vacuolaria with
discoidal chromatophores of a yellow-green colour, but more
strictly a Flagellate. Great interest is likewise attached to the
discovery by Luther in 1898 of yet another similar organism
3

which he named Chlorosaccus. This organism has certain resem-

blances to Tetraspora, but is of a yellow-green colour with several
parietal disc-like chromatophores, and seems
to connect Chlo-
ramoeba and Vacuolaria with the direct line of descent of the
Confervales. Luther proposed to remove all these forms out of
'

the Chlorophycese and suggested the name Heterokontce as a

'

class equal with that of the Chlorophycese, and to include the

'

Algal series
'
Confervales and the corresponding Flagellate group
'

Chloromonadina Chloromonadales). This class seems a very

'

(or
natural one and differs from the Chlorophycese in certain cyto-

logical characters, such as the abundant presence of xanthophyll

and the presence of a fatty substance as the stored product of
carbon-assimilation.
Bohlin has recently suggested that the Vaucheriacese should
' '

be included with the Confervales and Chloromonadales as one

' '

of the orders of the Heterokontce, with the name Vaucheriales.'

'

There are, however, wide differences in structure between the

Vaucheriacece and the Confervales, not to mention the absence of
the yellow pigment and the highly differentiated sexual repro-
duction present in the former neither is there any evidence that
;

the plants of these orders are in any way phylogenetically related ;

in
therefore, for the present, I prefer to retain the Vaucheriacese
the order Siphonea- of the Green Algae.
The origin of the Phseophycese, or Brown Algas, from brown

1
Cienkowski in Archiv. fiir Mikroscop. Anat. vi, 1867.
-
Bohlin in'Ofvers. af K. Vet.-Akad. Forh. 1897, no. 9.
3
Luther in Biliang till K. Sv. Vet.-Akad. Handl. Bd xxiv, 1898, no. 13.
30 Algae
 Phylogeini 31

Flagellate organisms seems quite as reasonable as in the parallel

case of the Green Algae. During recent years many genera of
primitive brown Algae have been discovered, most of which appear
to be intermediate forms between the higher brown Flagellates and
the simpler types of filamentous brown Algse. The majority of
these primitive brown Algae inhabit fresh water, but few of them
have up to the present been observed in Britain. The Flagellate
organism suggested as the possible starting point for this series is
Chromulina Cienk. 1 and the ascending series of forms include
,

Phosocystis Lagerh., Phceosphcera West & G. S. West, Phceococcus

Borzi, Entodesmis Borzi, Pkceothamnion Lagerh. and Pleurocladia
A. Br. Divergences along other lines from Chromulina may have
given rise to the Hydruraceae, the Chrysomonadinaceae and the
Dinobryaceae. PhcBodactylon Bohlin and Stichoglcea Chodat may
have a relationship with the Phceococcus-type of brown Alga.

PILEOPHYCE.E
A

Pleurocladia

PhjBothamnion

Entodesmis
Phaeococcus Phaeodactyloii
Stichogloea

Dinobryacete

Phaeosphaera
Hydruracese Phsocystis
Chrysoinonadaceiu

Chromuliua
PEOTOMASTiaiXA

The origin of the Bacillariese is still extremely doubtful, and

no reasonable suggestions have yet been put forward as to their

;
Cf. Lagerheim in Ofvers. af K. Vet.-Akad. Forb. 1896, no. 4, p. 288.
32 Algce
line of descent. They are regarded by some as a group of the

Phseophyceae, but are better considered as a distinct class. It

may be that there is a much more direct relationship between the

Bacillariea? and the Flagellate Peridiniese than is at first
apparent.
The origin of the large class Rhodophycea?
is still very un-

certain, although quite recently a marine Flagellate with the

chromatophores of the Rhodophyceas has been discovered by
Karsten 1 As there are so few freshwater representatives of this
.

large class of Algae, a discussion as to their origin would here be

out of place.
So little is known concerning the Myxophyceas and their life-
histories that any attempt to give an account of their origin and
evolution would be mostly a matter of conjecture. It is interesting
to note, however, the existence of blue-green motile organisms
such as Cryptoglena Ehrenb. and Chroomonas Hansg. 2 Reasons
'

for retaining the word

'

Myxophycea? are stated in the Intro-

duction (page 3).
The classification commonly adopted at the present time is the
Engler and Prantl's Pflanzenfamilien,' in which
'
one found in
the Green Alga? were arranged by Wille, the Bacillariea? by Schtitt,
and the Blue-green Alga? by Kirchner. Wille followed Sachs in
the removal of the Conjugate from the Chlorophycea?, but the
reasons for this are certainly insufficient. It is also significant to
note that since the publication of the classification referred to,
Wille has regarded the Conjugate as an order of Chlorophyceas 3 .

The most recent classification of Green Alga? in English is that

put forward by Blackmail and Tansley in the New Phytologist
4 ' '

for 1902, and they not only accept at the outset the
principal
changes suggested by Luther and Bohlin, but carry them still
further. They separate the CEdogoniales (as the "Stephanokontse")
and the Conjugate (as the "Akontae") from all the rest of the
"
Chlorophycea?, which are placed under the Isokonta3." This
arrangement is based upon the assumption that the CEdogoniales
and the Conjugate are phylogenetically independent of the
"
Isokontse," and that all three groups have arisen from the
1
Karsten in Wissensch. Meeresuntersuchungen, Kiel, Bd iii, Heft 2, 1898.
-
Hansgirg, 'Noch einmal liber die Phykochromaceen-Sch warmer,' Bot. Cen-
tralbl.Bd xxiv, 1885.
3
Wille. Algologische Notizen vn, vni,' Nyt Magazin f. Naturvideusk., B. 39,
'

H. 1, Kristiania, 1901.
4
F. F. Blackman and A. G. Tansley, 'A Revision of the Classification of the
Green Algae,' The New Phytologist, 1902.
 Phytogeny
Flagellata. Be
remembered, however, that there is no
it

evidence in of the view that the GEdogoniales ;ml

support
Conjugatoe are phylogenetically independent of the rest of the
Chlorophyceae. Indeed, with regard to the Conjugatse all the
known facts concerning them tend to show that they, at least,
have not had a direct origin from Flagellate ancestors. I have tin-
many years made a special study of the Conjugate?, including a
particularly detailed investigation of the Desmidiacea the family
11
,

around which all the interest of this idea is centred and I can ;

say most emphatically that all the facts concerning these plants
with which I am acquainted, far from supporting the idea of a
Flagellate ancestry, tend to prove conclusively that this beautiful
family of Conjugates has originated from filamentous ancestors.
The separation of the Conjugate from the rest of the
Chlorophycea3 is therefore to my mind based upon an erroneous
supposition, and there is likewise no evidence to show that the
complete separation of the (Edogoniales from the rest of the green
Algae is a just one. The Heterokontse is obviously a very natural
but I have not transferred the Vaucheriacese to the Hetero-
class,
kontas nor the Cladophoracea? to the Siphonese. The. genus
Prasiola has no relationship to the Ulotrichaceae and I have
placed it the order Schizogoniales. My arrangement of the
in

Conjugate also quite different from that given by Blackman

is

and Tansley, particularly in the family Desmidiacea?, of which I

have given a natural classification.
In those facts lie the main differences between the classification
put forward by Blackman and Tansley and the one used for the
green Alga- in the present volume, which is based upon a very
extensive and careful study of these plants for many years. There
will also be found many differences in the genera themselves and
in their disposition, as I have arranged them in a manner which I
consider to be much more in accordance with their affinities. For
example, the genera Radio/Hum Schmidle and Hormospora Breb.
are undoubtedly feebly developed forms of the Ulotrichaceae and
have no place in the Protococcacese and the same is true of the
;

genera and Stichococcus Nag. Dacti/lothece Hagerh.

Gloeotila Klitz.
is an
elongated Glceocystis-like genus and has no relationship with
either Dimorphococcus A. Br. or Scenedesmus Meyen and many ;

other instances could be quoted.

w. A.
 Class 1. KHODOPHYCE^; (or Floridese).

THIS class comprises the Algse usually known as the Red

Seaweeds. They exhibit a variety of colours from bright red and
purple to dark brownish-reds, brownish-green, blue-green and
black. Very few of the genera are freshwater forms.
The thallus is multicellular, very diverse in form, and consists
of simple or branched cell-filaments which may be merely held in
close proximity by mucilaginous material, or bound together by a

tough intercellular substance, giving the thallus a parenchymatous

appearance. The branching of the cell-filaments is very varied,
the plants exhibiting many types of branching. The filaments
increase in length by the repeated divisions of an apical cell.
The cells are all protoplasmically continuous through pits in the

transverse or cross-walls of the filaments. Each cell may contain

one well differentiated nucleus, or more rarely the thallus may
consist largely of coenocytes. The chromatophores are distinct
and the chlorophyll is masked by either a red colouring-matter
phycoerythriri, or a blue colouring-matter phycocyanin. Some-
times pyrenoids are present.
Asexual reproduction takes place by motionless spores known
as tetraspores (or tetragonidia), which are generally bright red,
and have neither cilia nor cell-wall. They are produced in a

tetrasporangium (or tetragonidangium), usually in variously ar-

ranged groups of four. This is the most common form of repro-
duction. Zoogonidia are absent from the entire class.
Sexual reproduction by male and female cells is wanting in
some, but present in others. The female organ is a procarp
which consists of a single cell containing a cell-nucleus, the
carpogonium, drawn out into an attenuated, hair-like process called
the trichogyne. The latter is homologous with the receptive spot
 Hhodophycece 35

of the oosphere of the green Alga*. The male organ is an an-

theridium of variable form which gives origin to large numbers of
male cells. Each male cell opens at the apex and sets free a
rounded, nucleated mass of protoplasm, without a cell-wall and
without cilia, known as a spermatium (or pollinoid). Fertilization
takes place by the attachment of the spermatium to the apex of
the trichogyne and the union of their contents. As the nucleus
of the spermatium disappears, it travels down the trichogyne and
unites with that of the carpogonium, this fusion having been
observed by Osterhaut in Batrachoxpermum Boryanum 1
.

The result of the fertilization of the carpogonium is the

development of a fructification known as a cystocarp (or sporocarp),
and the different groups of the Rhoclophycese are characterized by
the method of formation of this fructification. Sometimes the
cystocarp developed directly from the carpogonium but, fre-
is ;

quently, the fertilizing influence is handed on to other cells

in
the neighbourhood of the carpogonium, and conjugation occurs
between outgrowths of the fertilized carpogonium (known as
ooblastem a filaments) and certain auxiliary cells, the final result in
all cases being the development of the cystocarp and the production

of carpospores. Whatever be the method of formation of the

cystocarp, the carpospores always developed on a tuft of

are
filaments which spring from fertilized cells and which are known

as goniinoblasts.
The class is subdivided into four orders :

Order I. Nemalionacece. This order includes four families,

of which the Lemaneacese is exclusively fresh-
water, and the Helminthocladiea? includes several
freshwater genera.

Order II. Cryptonemiacew. One

family of this order, the
of which there
Squamariacese, contains a genus
are several freshwater species.

Order III. Gigartinacece. Exclusively marine.

Order IV. Rhodymeniacece. Exclusively marine.
In addition to the four orders just enumerated, another group
of Algae known as the Bangiacew is often included in the Rhodo-

phycece, but the systematic position

of this group is very uncertain.

1
Osterhaut in Flora, Ixxxvii, 1900.
32
36 Rhodophycece
The main argument for its inclusion in the Rhodophycese is

derived from the red colour of the chromatophores, whereas the

intercalary growth of
the thallus and the absence of the pits
between the thallus-cells are points against its inclusion among
the Red so-called tetraspores of the Bangiaces? are
The
Algse.
somewhat remarkable, the whole of the contents of a thallus-cell
amcebiform spore. The sexual
going to form one unciliated,
is also of a very reduced type, far removed even
from that
process
of the simplest red Alga. The genus Bangia, which is a simple

filamentous form, occurs on the shores of the British Islands and

in the estuaries of the rivers, but it is not exactly a freshwater Alga.

Order I. NEMALIONACE^E.
The fertilized carpogonium gives origin directly to the gonimo-
blasts, which are developed in tufts.

Family 1. HELMINTHOCLADIEJE.
The thallus
is filamentous, simple or branched, with the
axes often arranged in whorls. The main axis may
secondary
consist of a single row of cells, or of a central cell-filament

surrounded by a cortical ring of smaller cell-filaments. The

of cell-filaments and the terminal cells
gonimoblast is a short tuft
usually form the carpospores.
When the terminal cell has shed
its carpospore, the supporting cell grows through into the old cell-

wall and produces a new spore-forming cell. There is no definite

wall to the cystocarp.
Genus Batrachospermum Roth, 17!>7. This an exclusively
is

freshwater genus with a wide distribution intemperate and

Most of the species prefer deep water in
which
tropical climates
there a slight current, but more rarely they are found attached
is

to stones in fast streams. They scarcely ever occur in stagnant

water, but are found frequently in bogs, usually at a point
where
a spring rises. The thallus, which is of a blue-green colour and
in a thick coat of mucus, is remarkable for the great
enveloped
it reaches a
beauty and symmetry of its branching; sometimes
length of 16 20 cms. The plants are generally attached to
 Heltt t it ithocladiece 37

wood by a number of thick

stones or old shoots ;
these send off
numerous primary axes which float freely in the water. Tin-

primary axis consists of a central filament of cells which grows by

Fig. 1. A, Batrachospeninan monilit'orine Both, from Malham Cove, W. Yorks. ( x 2);

B C, Batrachospermum vagum (Both) Ag., from Thursley Common, Surrey:
B (
x 2) C, single node with lateral branches, more highly magnified. D, ger-
;

minating spore. E, protonemal growth. F, Female organ and fertilization;

s, spermatium; c, carpogonium ; t, trichogyne. (D, E, and F after Schmidle.)

means of a hemispherical apical cell. The cells of this central

filament become swollen at each end, a dense whorl of branches

being produced at each swelling (or node).

From the basal cells
38 RhodopTiycecB

of the branches secondary branches grow downwards over the

main axis, forming a cortical ring of cell-filaments (sometimes
termed the pseudocortex). The apical cells of the lateral branches
are frequently produced out into long hairs or bristles. The pro-
is unicellular and is developed at the extremity of a small
carp
branch which stands out directly from the main axis, and which is
termed by Sirodot the female axis.' The carpogonium possesses
'

a short, straight, exposed trichogyne and after fertilization it

1
,

develops the dense mass of gonimoblasts (frequently termed a

glomerulus ') which constitute the cystocarp. The cystocarps are
'

external, being developed amongst the dense whorls of branches,

and the terminal cell of each gonimoblast produces a carpospore.

On the development of the carpospores sporophytic shoots are

formed which very much resemble species of the genus Chan-
trim .via: they are to be regarded as a kind of 'protonema,' which,
under certain suitable conditions, develops into the sexual Batra-
cli
osperrait w-plant.

There are two more or less abundant species of the genus in the British
Islands, B. moniliforme Roth (Fig. 1 A) and B. vet gum (Roth) Ag. (Fig. 1 B),
each with a number of varieties. The latter is distinguished from the former
by the more or less uniform development of the lateral branches along both
the nodes and internodes of the inferior portions of the primary axis. A
third species, B. atrum (Dillw.) Harv., with very short lateral branches and
long internodes, is less widely distributed. Species of this genus commonly
afford a home for epiphytes of the nature of certain species of Calothrix,

Hapalosiphon, Ammatoidea, etc., and numerous Diatoms and Desmids are

frequently present in their enveloping mucus.

Fries, 1825. The plants belonging to this

Genus Chantransia
genus are much
smaller than species of Batrachospermum and
occur both in fresh water and in the sea. The freshwater species
vary from about 3 to 7 mm. in length, and are usually found
attached to rocks and stones in rapid rivers, cataracts and water-
falls. The thallus consists of branched filaments of cells arising
from a basal stratum, the apices of the branches being frequently
much attenuated and almost piliferous. In colour the plants are
all shades of red, purple and blue. There is an entire absence of
the mucous coat which is so conspicuous a feature of Batracho-

spermum. It was thought for some time that all the species of

Schmidle, 'Einiges iiber die Befruchtung, Keimung, und Haarinsertion von

1

Batrachospermum,' Bot. Zeitung, Heft 7, 1899.

 Helininthocladie j

Chantransia were merely non-sexual stages of Batrachospermum,

and that under brighter condi-
tions of light they underwent a

metamorphosis, giving rise to

the sexual stage or Batracho-
spermum. The carpospores of
the latter genus develop a pro-
tonema-like growth which bears
of
great resemblance to species
Chantransia, and many of these
growths, both of Batrachosper-
mum and Lemanea, have been
erroneously described as species
of that genus but, at the same
;

time, these have nothing to do

with the valid genus Chantransia.
The sexual reproduction of Chan-
transia has only been fully worked
out in one species Ch. coryin-
bifera Thur. On the fertilization
of the carpogonium it develops
numerous gonimoblasts upward-
ly and on one side. There
is thus

formed a naked corymbose cysto-

carp, the terminal cells of the

gonimoblasts producing the car-

pospores. The antheridia are
likewise developed in clusters.
Asexual reproduction occurs by
tetraspores and also by other
spores which remain undivided
and are known as
'
Fig. 2. A, CJiiuitniiixiii pii<i>n<r<i
monospores.'
Kiitz., from Penyghent, W. Vurks.
These, on germination, divide (
x 100). B and C, branches of the
same with 'monospores' (m) x 300).
into four cells in one plane,
(

D, Ch. scotica Kiitz., from Cornwall a :

giving rise to the basal stratum small portion of the thallus showing
from which the branched fila- the pits in the transverse walls ( x 400).

ments spring.

There are some seven or eight British freshwater species of the genus, <>f
which Ch. pygmtm Kiitz. (fig. 2 A C) and Ch. violacca Kiitz. are perhaps the
most frequent.
40 RhodophycecB
Genus Thorea Bory, 1808. This is a rare genus with only
one species ramosissima Bory which, although found in
Th.
several of the rivers of France and Germany, has only once been
recorded for Britain (from Walton-on-Thames). It has a round
filamentous thallus, which is much branched and reaches a length
of 30 60 cms. It is about the thickness of a horse-hai r ,
of a

purple-brown or dark brown

colour, and very mucous. There is a
central solid axis consisting of filaments of cells, and arising from
this axis are a very large number of short compact branches, which
are slightly attenuated. The cells of the branches are from two
to five times longer than their diameter. Schmidle has recently 1

worked out much of its structure and fructification, and this has
been largely confirmed by Hedgcock and Hunter 2
This genus has .

been placed under the Phseophycese, but the pigment, the presence
of starch-like granules in the cells, and the naked non-motile

spores, indicate a
near relationship with certain of the Floridea?.

Family 2. LEMANEACE.S3.
This is a small group of exclusively freshwater Algae including
the two genera Lemanea and Sackeria. They are plants which
only grow in rapid torrents, occurring attached to the rocks of
waterfalls, to stones and wood in mill-sluices, etc., always where
the force of the water is greatest. The thallus is composed of a
3 "
basal, attached portion, termed by Sirodot a systeme radicant,"
from which arise csespitose tufts of erect, branched, simple fila-
ments reaching a length of 3 8 mm. From portions of these
filaments the fructiferous branches These are the most
arise.

conspicuous and important parts of the plant, in most species the

vegetative portion dying away after their production, and in a short
time they become fixed by organs of attachment of their own.
Each species is thus represented by two distinct sets of indi-

viduals, the one vegetative and the other reproductive.

The fructiferous branches are elongated, thread-like portions
1
Schmidle, Uutersuchungen iiber Thorea ramosissima Bory,' Hedwigia,
'
Bd
xxxv, 1896.
2
G. G. Hedgcock and A. A. Hunter in Botan. Gazette, xxxviii, 1899.
3
Sirodot,
'
Etude anatomique, organogenique, et physiologique de la Fam. des
Lemaneacees,' Ann. Sci. Nat. Bot. torn, xvi, Paris, 1872.
 Lemaneacece 41

of the thallus, of a cartilaginous

consistency, and hang freely in
the rapid torrent. They are of an olive-green or greenish-Mirk
colour and grow to a length of 15 18 cms. At short, inoiv or

Fig. 3. A, Saclu'i-in mamillosa Sirodot (nat. size), from E. Wharfe, W. Yorks.

B, portion slightly magnified showing antheridial areas. C, Lcmmifn tnnilomi
Kiitz. (nat. size). D, portion slightly magnified showing antheridial areas.
E, Suclierin fucina (Bory) Sirodot, longitudinal section of fructiferous filament
showing female organ t, trichogyne. F, Lcinnin'ii cuicnata Kiitz., transverse
;

section of fructiferous filaments showing antheridia; a, antheridial cell;

, spermatium. (D, E, and F after Sirodot.)

less regular intervals along their whole length are distinct swell-
ings or nodes, and each nodulose thread is built up of an axial
42 RTiodophycece
row of tubular cells surrounded by rows of smaller cortical cells,
growth taking place in all cases by an apical cell. The only
known method of reproduction is a sexual one. The antheridia
are short, cylindrical cells developed on the exterior of the thallus,
either on verticillate eminences or on the widest parts of the
nodes. The procarp is unicellular and the carpogonium possesses
a rather long, transparent, simple or branched trichogyne. After
the fertilization of the carpogonium an ooblastema-filament is

developed from it, at the extremity of which a bunch of jointed,

moniliform filaments arise. Each of the swollen cells of these
moniliform filaments becomes, when mature, a carpospore.
The carpospores are thus produced inside the thallus, filling

up the space between the central axis and the cortical cells
of the fructiferous On development the carpospores.
filaments.

produce the vegetative thallus, protonema-like growth which has

a

frequently been mistaken for a form of Ckantransia.

Genus Lemanea Bory, 1808. The vegetative thallus generally
persists for several months, and is very branched but never pili-
ferous. The fructiferous filaments are torulose, being regularly
inflated at even distances, and are normally simple. The central
axis of tubular cells is surrounded by a series of spirally twisted

axial filaments,and the antheridial areas are in more or less com-

plete rings round the nodes.
L. torulosa Kiitz. ;
em. Sirodot (fig. 3 C, D), and L. parvula Sirodot, are
found in the rapid streams and torrents of certain parts of the British Isles.

Genus Sacheria Sirodot, 1872. The vegetative thallus is very

fugacious. It is little branched, often piliferous, and exists for
about a month (generally December or January). The fructiferous
filaments are cylindrical or setaceous and usually branched. The
central axis of tubular cells is
quite devoid of axial filaments and
the^ antheridial areas are on mamilliform projections, arranged in
a verticillate manner at regular intervals.

Species of this genus are much more frequent than species of Lemanea*
Three species of the genus are widely distributed in the British Isles,
S. fluviatilis (Ag.) Sirod. (syn. Lemanea fluviatilis Ag.), S. fucina (Bory)

Sirod., and S. mamillosa Sirod. (fig. 3 A, B), the last-mentioned one being
the most abundant. It appears that >V. mamillosa may sometimes occur in
still water, as I have recently examined specimens of this species collected by

Mr J. Murray from the west side of Loch Ness, Inverness. It must be

remembered, however, that Loch Ness is a large body of water forming part
of the Caledonian Canal, and probably numerous currents exist in it.
 Squamariacece

Order II. CRYPTO NEMIACE^.

The fertilized carpogonium sends out a long, branched oobh--
tema-filarnent, the terminal cells of each branch conjugating with
an auxiliary cell. From the latter the gonimoblasts arise.

Family 1. SQUAMARIACE.E.
This family consists of a small group of marine, or rarely fresh-
water Algae, which are minute, flat, gelatinous or membranous
expansions, commonly encrusting stones,
shells, or larger Algae. The thallus usually
consists of dense, vertically arranged cell-
filaments. Tetrasporangia are formed in
various ways, and often give the surface of
the thallus quite a verruculate appearance.
The sexual organs are developed in cavities
'

conceptacles in the upper surface of the

'
or
thallus. After the fertilization of the carpo-
'

gonia these conceptacles contain numerous

'

Fig. 4. Hildenbrandt-
cystocarps. ia rivulari* (Liebin.) J.
Genus Hildenbrandtia Nardo, 1845. Ag., from Shipley Glen,
W. Yorks. A, section
This genus consists of a crustaceous expanded of thallus. B, surface
thallus, of a blood-red, dark red, rose, or view (x400). C, two
cells showing the chro-
brown colour, firmly adhering to rocks and
matophores ( x 800).
stones on the sea-shore or in rivers and
streams. The thallus is composed of compact, vertically arranged
cell-filaments, with subcubical or oblong cells. The cell-walls are
colourless and strong.

There is only one British freshwater species H. ri>->il<i ,-!.< (Liebin.) .T. A-,
(fig. 4) which occurs as dark red patches on rocks and stones in streams and
dripping places.
 Class 2. PH^OPHYCE^E (or Fucoidese).

ALMOST all the Algse of this class are marine and are known as
theBrown Seaweeds. They are often termed the Melanophycese.
The thallus exhibits great diversity of form in some it is a ;

simple filament, in others a flat expansion of cells, and in others it

is
greatly differentiated. The most highly organized of all seaweeds
are members of this class of the brown Algse. The vegetative
cells possess one nucleus, and the chromatophores have a distinct
brown tinge owing to the presence of phycophcein and phycoxanthw.
(the compound pigment being known as phaeophyll). The former
can be extracted with water and the latter by means of alcohol.
Asexual reproduction is by motile cells or zoogonidia.
Sexual reproduction is either by isogamous or heterogamous
gametes, the conjugation of the gametes or the fertilization of the
oospheres taking place in all cases outside the plant. The zygospore
or the oospore always germinates directly. The motile reproductive
cells,whether zoogonidia or gametes, invariably possess two cilia,
inserted laterally, and in their movements one cilium is carried in
a forward direction and one in a backward direction.
The class is divided into a number of orders of which only one
the Syngeneticcv is freshwater.

Order I. SYNGENETIC^E.
The plants included in this order, which is sometimes termed
'
the Phasozoosporina?,' are exclusively freshwater. They are Alg*
of little note or importance, and may be either solitary or colonial
unicells, multicellular, free-swimming or motionless. The cells are

often naked, but at other times are surrounded by a mucilaginous

cell-wall. In some of the multicellular forms the cells are only
loosely held in position by a copious mucilaginous envelope.
There
is one cell-nucleus and one or more pulsating vacuoles. The
 Hydruracece 4.">

chromatophores, which are of a yellow or pale brown colour. inn\

be solitary and excentric, or sometimes two in a cell, and pyrenoids
are occasionally present. Vegetative multiplication occurs by
simple cell-fission and asexual reproduction takes place by me;m^
of zoogonidia.

The following seven families are the most important :

Fam. Hydruracea'. Hi/<lnirns Ag.

Fam. Cryptomonadinacece. Cryptomonas Ehrenb.
Fam. Ghrysomonadiiwcece. Xyttcrypta Ehrenb. Syntinr
Ehrenb. Uroylena Ehrenb.
Fam. Dinobryacecs. Dinobryon Ehrenb.
Fam. Phwocapsacece. Plumcystis Lagerh. Pltceococcus
Borzi. Phceosphcera West & G. S. West.
Stichoyloea Chodat. Phceschizochlamys Lemm.
PJueodactylon Bohlin.
Fam. Choristocarpacece. Plenrocladia A. Br.
Fam. Phrvotliamniacece. Phceothamnion Lagerh.

Only four of the above families are known to be represented

in Britain, but probably all the others will be found in more or less
abundance if searched for. Many of the plants of this order are
plankton-forms, and the families Cryptomonadinacese, Chryso-
moriadinaceff' and Dinobryacese are truly flagellate in character.

Family 1. HYDRURACE^I.
The plants are attached, branched, and consist of a colony of
unicells. The but afterwards become
cells are at first spherical

almost spindle-shaped, and are embedded in large masses of

mucilage. They have one chromatophore and are destitute of a
cell-wall. Asexual takes place by zoogonidia which
reproduction
are tetrahedric in form and possess one cilium
1
.
They are only

produced in the branches, two or four from each cell, and they
germinate directly. After having come to rest, the zoogonidia
attach themselves by the clear apex (at which point the cilium
was inserted) and secrete a stalk-like mass of mucilage. This cell
is the
beginning of a new colony which is developed subsequently
take on
by its repeated divisions. Certain of the peripheral cells
apical growth and produce branches. Resting spores (akinetes)
have also been observed by Lagerheim.
1
Cfr Lagerheim in Ber. Deutsch. Bot. Gesell. 1888, p. 80, tig. xylogr. 13.
46 Phceophycece
Genus Hydrurus Ag., 1824. The plants are branched colonies
of unicellular units embedded in a tough, cylindrical mucilage.

They vary from about 5 to 30 cms. in length

and are of an olive green colour. The
whole colony is simple below but branched
above, often cut up into fine penicillate
divisions, and covered with small villous
projections giving it a plumose appearance.
The entire structure behaves almost as a
multicellular plant, growth in length being

entirely dependent on single apical cells,

and the branching is usually monopodial.
& The cells are commonly ellipsoidal and are
more densely crowded in the small branches
Fig. 5. Hi/drums than in the main stems and branches. After-
Kirchn.
fcetidus (Vill.) wards the cells elongate and become ar-
A, nat. size. B, zoogoui-
clium (after Lagerheim). ranged more or less, in longitudinal groups.
H. fcetidus (Vill.) Kirchn. is found attached to stones and rocks in
mountain streams. It is a sticky plant and gives off an offensive odour when
alive. It is common in Central Europe and in the Arctic regions when the
snows melt in the spring, but in the British Islands it is of very rare occur-
rence, being known only from Yorkshire and Scotland.

Family 2. CHRYSOMONADINACE^.
These are unicellular or colonial organisms which in the free
condition are motile. Each individual
consists of an oval or elongated cell, with
either one or two cilia and either one or
two brownish-green chromatophores. A
red pigment spot is generally visible. The
cells increase by longitudinal division.
Genus Synura Ehrenb., 1838. This is
a small, globose, free-swimming colony,
Fig. 6. Synura Uvella formed of a variable number (from 10 to
Ehrenl). Single colony
(x400), from Eldwick, 50) of ovoid or ellipsoid, biciliated indi-
W. Yorks.
They are arranged close together
viduals.
in a radialmanner, and each individual possesses two chromato-
phores and at the hinder end two pulsating vacuoles.
commonly found in small ditches and
Synura Uvella Ehrenb. (fig. 6) is

pools, particularly they are of rain-water.

if Pure collections of it can be
frequently obtained in the early summer.
 Dinobryacew 47

Genus Syncrypta Ehrenb., 1838. This is a motile colony

similar in appearance to Synura but invested with a mucilaginous
coat through which the cilia protrude.

Syncrypta Volvox Ehrenb. is an abundant organism which bears great

resemblance to Synurct Uctfla.

Genus Uroglena Ehrenb., 1838. In this colonial form tin-

cells are of the same nature as those of Synura, but the central
is a hollow space filled with mucilage, and
portion of the colony
the ciliated cells are arranged round the periphery.

Volcox Ehrenb. is found in similar situations to those mentioned

t/

for the two previous genera. It is, however, much less abundant.

Family 3. DINOBRYACE.E.
The individuals are attached to the bottom of a cup-shaped

receptacle, which is widely

are con-
open above. They
tractileand possess two cilia
of unequal length.
Genus Dinobryon Ehr.,
1833. The cells are very
delicate, of a somewhat
changeable form, and are
sensitive to stimuli. The
lower end is attenuated into
a stalk which is attached
near the base of the open
receptacle. There is one
long cilium and one shorter
secondary cilium. The chro-
matophores are two in num-
ber and of a yellow -brown
colour. There is a pigment
spot,two contractile vacuoles, Fig. 7. A, Dinobryon cyliiuJricum Iinhof
Lemm.; two
var. dirertjcnit living examples
and one cell-nucleus. The from Eldwick, W. Yorks. x 730). B, en-
(

cysted condition same.

receptacle is campanulate or of C, Dinnliri/nn
S/Ttularia- Ehrenb., colony with individuals
cylindrical, attenuated at its from Cornwall x410); c, cysts.
encysted, (

lower end into a straight or

oblique point; it is hyaline or sometimes coloured yellow or bn>\vn
48 Phceophycece
with oxide of iron, and the margins may be smooth or undulate.
The by longitudinal division or by the formation
multiplication is

of globose resting-cells (or cysts) which are furnished with a

peculiar projecting process (fig. 7 B and C). The cells occur singly
or joined into dense, spreading colonies. The daughter-cells effect
a lodgement above the inner rirn of the mother-receptacle and
then secrete a similar receptacle for themselves. Senn has written 1

a good account of this genus, and Lemmermann' has published a 2

monograph of it, discriminating between fourteen species.

Three species, D. Sertularia Ehrenb. (tig. 7 C), D. sociale Ehrenb. and
I).
I'yUndricum Imhof, and varieties of them, are abundant throughout the
British Isles, the first-named one being the most widely distributed.
Species
of this genus are very abundant in the freshwater plankton, the colonies of
each species exhibiting a characteristic type of branching. D. protuberans
Lcmm. and D. elongatum Imhof are generally distributed but not abundant.

Family 4. PILE] OC APS ACE^E.

The plants are unicellular, forming colonies, the cells of which

are embedded in a mass of mucilage. The cells are spherical or
ellipsoidal and division takes place in all directions. The repro-
duction by zoogonidia and zoogametes.
is

Genus Phaeococcus Borzi, 1892 The cells are :i

.
ellipsoidal or
oblong-ellipsoidal, 6 llyu, in diameter,
and occur in twos, fours, eights or mul-

tiples of these numbers, in hyaline

r*
Qf) . gelatinous integuments which some-
A times show a delicate concentric struc-
ture. There are two yellow-brown
chromatophores in each cell and usually
a red pigment-spot. The zoogonidia
are ovoid or subpyriform.
tig. 8. Phccococcus palu-
dosux West & Or. S. West, from
Eldwick, W. Yorka. x 410). z,
P ( lementi (Menegh.) Bora has not been
'
/^ 7 . /-. r

zoogonidia. observed from Britain, but P. paludosus West

& G. S. West occurs in moorland ditches.

1
Senn, 'Flagellata' in Engler and Prantl Natiirl. Pflanzenfam. I Theil. la Abth.
2
Lemmermann, in Berichte Deutsch. Botan. Gesellsch. 1900, Bd xviii, pp. 500
524, t. xvii u. xix.
;i
Borzi in Atti del Congr. Botan. Internaz. Genova, 1892, pp. 463471, t. xviii.
 Phceocapsacete 49

Genus Phaeosphaera West & G. S. West, 1902. The cells

are large, exactly spherical, 14 17'5/i
in diameter, and are embedded in small

aggregates in a cylindrical, gelatinous

integument which is sparsely branched.
One brown, parietal chromatophore with
somewhat irregular margins is
present
in each cell.

P. gelatinosa West & G. S. West (fig. 9) is

known from Sphagnum-bogs in Cornwall.

Genus Stichogloea Chodat, 1897 1

.

The cells are small, oblong or subovoid

in shape, and are associated to form a

membranous, gelatinous thallus of small

size. The thallus is generally variously
lobed and the cells are often somewhat
radiately disposed. The cell-walls are
firm and each cell contains a
parietal
chromatophore destitute of a pyrenoid.
Fig. 9. Phaospluera gelati-
S. olivacea Chodat is known from the nosa West & G. S. West. A,
plankton of certain of the Scottish lakes. portion of colony (x50). B
and C, cells showing the solitary
Length of cells 9 15/t.
chromatophores ( x 410). From
Treniethick Moor, Cornwall.

1
Chodat in Bull. L'Herb. Boiss. torn, v, no. 4, 1897, p. 302, t. f. 812.
10,

W. A. 4
 Class 3. CHLOROPHYCE^E.
THIS group, which includes all the green Alga?, attains its
greatest development in fresh water, and the number of species
exceeds the combined total of the freshwater species of all other
Algae.
The simpler forms of green Alga? are unicellular (e.g. some of
the Protococcoideae and Desmidiacese), some are coenocytic (e.g.
Yaucherfacea3, Sphasropleacese, Pediastrea?), some are incompletely
septate (e.g. Cladophoracea?), and others are
multicellular or com-

pletely septate (e.g. CEdogoniales, Chtetophorales, Zygnemacese).

In other than the unicellular forms the thallus exhibits every
degree of development from simple rounded cells to long, simple
or branched filaments, flat expansions, or pulvinate masses of
tissue. As a rule there is no differentiation of the ordinary vegeta-
tive cells, but in some there is a marked distinction between the
vegetative and reproductive cells.

The cell-protoplasm (or cytoplasm} of the green Alga consists of 1

'
a lining layer or primordial utricle which adheres closely to the
'

cell- wall 1 and, in many Alga?, of additional anastomosing strands and

,

threads traversing the interior of the cell. It contains numerous

granules of variable size which behave differently with staining
reagents. Evidence goes to prove that there is no definite proto-
plasmic continuity between the cells of multicellular green Algae.
A division of labour is rarely observed amongst this class of plants,
and in the Conjugata? the cells of most of the filamentous forms
arc under normal circumstances quite able to lead an independent
existence. The vacuoles are much as in other plant-cells and they
contain a fluid usually known as the cell-sap. In the Conjugate
the cell-sap is occasionally coloured violet or purple owing to the

Cf. Chodat et Boubier,

1 '
Sur la Plasmolyse et la membrane plasmique,'
Journ. Bot. de Morot, Paris, 1898.
 Cfdorophyceeb 51

presence of a pigment termed by Lagerheim phycoporphyriri

1
.

This violet colour occurs normally in Ancylonema Nordenskioldii

Berggr., Mesotwnium violascens De Bary, M. purpureum W. &
G. S. West and Mougeotia capucina (Bory) Ag., and under excep-
tional circumstances it is found in various species of Zygnema,
Spirogyra and Desmids. The Volvocacege and the zoogonidia and
of other green Alga- possess vibratile cilia, which are very
1

gametes
variable in their length, number, disposition, and symmetry; and
in certain of the same forms contractile vacuoles are present. In
'
the genera Tetraspora and Apiocystis pseudocilia are found,
'

which do not possess any power of movement.

A is present in the cells of all the green
single nucleus Algae
except the coenocytic and incompletely septate forms, and during
the formation of asexual non-motile spores, zoogonidia, or gametes,
it
undergoes divisions corresponding to the divisions of the proto-
plasm. In some green Alga? mitotic division of a more or less
complex character has been observed-.
The and its structure is often difficult
cell-wall is very variable
of observation. In the formation of a cell-wall such as after the
quiescence of a zoogonidium, it is developed
on the outer surface
of the protoplasm as the result of more or less complex processes.
The young cell- wall usually consists of cellulose, but sometimes
equally of pectose. Under the action of strong acids or other
will swell up and
hydrating reagents an ordinary thick cell-wall
show traces of lamination. Each lamina represents successive
layers of growth in thickness and in
most plants consists of a
mixture of cellulose and pectose constituents in variable propor-
tions. In the Chlorophycese these two constituents of the cell-
wall are differentiated while the wall is very young. They exhibit
considerable differences in behaviour with reagents, the
their
cellulose constituents giving a violet colour with chlor-zinc-iodine

(Schulze's solution), whereas the pectose constituents do not. In

in the
many Algse the pectose constituents of the cell-wall are
form of gelatinous layers on the outside of inner layers of cellulose.
This mucilaginous material stains readily with aniline dyes such

1
Lagerheim in Videosk.-Selsk. Skrift., I niatbem.-uatur. Kl., Kristiania, 1895,
no. 5.
Observed in Spirogyra by Mit/kewitsch (Flora, Ixxxv, 1898) and C. van
2

Wisselinsh (Bot. Zeitung, Ivi, 1898; Flora, Ixxxvii, 1900); in Chlamydowonn* by

Dangeard (Le Botaniste, vi, 1899) in Closterium by Klebahn
;
also in llotry-
;

diurn, etc., etc.

42
52 Clilorophycem

as fuchsin, safranin, methylene-blue and gentian- violet. The outer

layers often become thick coats of mucilage by

the formation of
series of pectose constituents which exhibit all stages between
and complete solubility in water. It is not merely a
insolubility
hydration but a molecular change, and successive increments are
often added by the gelatinization of other layers of the cell-wall.
In some of the unicells the increase in thickness of the cell-wall
due to gelatinization is only on one side and elongated colonies
such as those of Hormotila are formed.
The gelatinous pectose compounds although sometimes forming
a large proportion of the cell -wall, do not alternate with layers of
but there appears to be a continual exudation of them
cellulose,

through the inner layers of cellulose, a mass of jelly being thus

formed on the outside of the cellulose wall. This is best seen in
some of the Protococcoidese and Conjugate. The mucilage in
which filaments of Algse are so frequently embedded exhibits a
distinct radiating fibrillar structure which is clearly brought out

by various reagents. The radiating structure of the enveloping

mucus has long been known in the Conjugates and has at times
given rise, especially in the Desmidiacea?, to grave morphological
misconceptions.
The (Edogonium and other Alga? exhibit peculiari-
cell-walls of
which will be described in their respective families.
ties of structure
Whatever the nature of the cell-wall one of its primary func-
tions is the regulation of osmotic changes.
Hairs and bristles are developed by certain green Algae belong-
ing to the Coleocha3tacea?, Herposteiracese, CEdogoniaceas, Chseto-
phoracese and ChastopeltidaB. They are of many kinds, from
slender articulate branches such as the piliferous apices of Chceto-
phora, to exceedingly fine inarticulate hairs such as the seta^ of
Bulbochcete, Herposteiron, ChcBtosphceridium, or Gonochcete.
The thallus often develops special root-like organs of attach-
ment or haptera (commonly termed rhizoids), but these are as a
rule only found in the young plants, most of the older ones occur-
ring as freely floating masses.
The chromatophores are usually distinct, and in the forms with
a filamentous or expanded thallus they are frequently characteristic
of the different families or genera. They are of a bright green
colour due to the presence of chlorophyll and are therefore cJiloro-
plasts. Sometimes they are very difficult to define, but at other
 Chlorophycece 53

times they stand out clearly, occupying only a relatively small

portion of the cytoplasm. They may be solitary or very numerous,
of infinite variety of form, central or parietal, and the edges may be
entire or deeply incised. In some forms they are ribbon-like and
wound round the interior of the cell-wall (e.g. Spirogyra,
spirally
Genicularia, and some species of Spirotcenia), and in others they
are central, spirally twisted masses (e.g. some species of Spirotcenia).
Sometimes they are reduced and very pale in colour, and in the
rhizoids and terminal cells of the hairs of some green Alga? they
are entirely wanting.
The
chloroplasts of most green Algae contain pyrenoids or
proteid bodies which serve as a reserve of food-material. Much
has been done towards the investigation of these bodies during
the past few years and the presence or absence of pyrenoids has
been regarded by some as a sufficient generic distinction. This is,
however, attaching an importance to these proteid bodies which is
scarcely borne out by facts. Although they frequently divide
equally on the division of the cell, they also multiply without any
cell-increase. They are likewise known to disappear during the
of certain and it has been clearly demon-
development species,
strated that during certain stages of Tetraspora, Spliceretta and
Eudorina they can arise spontaneously. Moreover, forms of An-
kistrodesmus falcatus (Corda) Ralfs containing pyrenoids are
sometimes met with in the same collection as others which have
no pyrenoids. Similarly, the chromatophores of Debarya calospora
(Palla) W. & G. S. West may or may not contain pyrenoids
1
.

Starvation causes a disappearance of pyrenoids and they frequently

increase in numbers if the cell is well nourished. On the whole,
there is doubt that the presence or absence of pyrenoids
little

depends largely upon external conditions and is a character to

which a great deal too much importance has been attached in
discriminating between the genera of green Algae.
In the Chlorophyceae the stored product of assimilation is almost
invariably starch. Exceptions to this are found commonly in
Mesotce mum in the Desmidiaceae and in the Vaucheriaceae.
in the of the
generally takes place
Cell-division all cells

but in a few instances there

thallus, is a definite growing point
which is usually an apical cell.

1
West & G. S. West in Ann. Bot. xx, March, 1898, p. 49; in Journ. Bot.
Aug. 1900, p. 289.
54 Chlorophycece

Multiplication by cell-division occurs in many of the lower

forms of the Protococcoidese and in the Desmidiacese. In the
Zygnemacece, and particularly in the smaller species of Spirogyra,
the filaments often dissociate into solitary cells each of which then
divides and forms a new filament.
Asexual reproduction by zoogonidia is general throughout the
class, although there is a notable exception in the Conjugata?, in
which motile reproductive cells are entirely wanting. In many of
the Chaetophorales, Microsporales and Protococcoidese reproduction
takes place by non-motile spores which may be either akinetes or
aplanospores. Asexual spores are also more rarely found in the
Conjugate, having been observed in Zyynema, Spirogyra, and in a
few Desmids, and they are formed normally in the rare genus
Gonatonenta.
Sexual reproduction occurs in most of the families of green
Algse, and may be either isogamous or heterogamous. In iso-
gamous reproduction the sexual organs are gametangia, usually

giving rise to planogametes which conjugate and produce a zygo-

spore. In the Conjugate only aplanogametes are found. Many
planogametes are generally produced from one gametangium, but
only one aplanogamete. In heterogamous reproduction the sexual
organs are oogonia and antheridia, and the gametes consist of
oospheres and antherozoids. In all cases with the exception of
Herposteiroii, the oosphere remains in situ in the parent plant,
being fertilized within the oogonium, and the result of fertilization
is an oospore. The oogonia are always unicellular, and, except in

Gylindrocapsa and some species of (Edogonium, so are the anthe-

ridia. Only one oosphere is produced in an oogonium, except in
SpJicvroplea, and one, two, or many antherozoids may arise from an
antheridium.
The gametophyte the principal generation, the sporophyte
is

being generally represented by the sexually-produced spore. In

Coleochwte, in (Edogonium, and in Mougeotia very rudimentary
sporophyte generations consisting of several cells do exist.
Of all freshwater Algse the Chlorophycese have the most varied
habitats. They are found in every possible damp or wet situation
and some are epiphytes, others endophytes, and a few are even
1
parasites .

1
PhyUosiphon Arisari Kiihn (in Sitzungsber. d. naturf. Ges. in Halle, 1878) is
a parasitic Alga observed only on the leaves of Arisarwn vulgare in Italy and the
 CMorophycece 55

Richter and Comere 2 have conducted experiments with a view

1

to ascertaining if certain of the freshwater Algse, especially Chloro-

phyceae, can exist in salt water. Richter states that the lower the
organization of the Alga the better its power of adaptation, but

Comere finds that only those Alga? with a robust structure and
Avith large chloroplasts can successfully withstand immersion in
salt water. Some species of (Edogonium and Cladophora can
live in Avater containing 3'5 / of sodium chloride, Vaucheria
water containing 2 / and some of the large species of
sessilis in ,

Spirogyra in water containing from T8 2/ Richter affirms .

that (Edogonium, Spirogyra, or Vaucheria have less power of

adaptation to life in salt Avater than Stichococcus or Tetraspora.
In all cases the salinity of the water caused the cells to increase in
size and Avhen the concentration was high malformation of the cells

invariably occurred. Starch at first disappeared from the cells,

but reappeared Avhen the adaptation Avas more complete. Not-
withstanding the somewhat contradictory nature of these two sets
of experiments, it appears that certain of the freshwater Chloro-

phycere can adapt themselves to an increasing salinity of the

water in a manner comparable with the adaptation of a feAV forms
of the green Algge to a life in hot water 3 .

The class Chlorophycese can be conveniently subdivided into

nine orders, all of Avhich are found abundantly in the British
Islands.

Order I. (Edogoniales. Thallus filamentous, simple or

branched. Cells uninucleate, Avith a large, parietal,

anastomosing chloroplast containing one or several

pyrenoids. Cell-division characterized by the
inter-
calation of a neAv piece of cell-Avail between the
mother-cell and the distal end of the daughter-cell.
Sexual reproduction by heterogamous gametes.
Zoogonidia Avith an anterior circle of cilia.
Ex-
clusively freshAvater.

south of France; P. maximus Lagerh., P. Philodendri Lagerh. and P.

Lagerh. are parasites on the leaves of species of Arisarum, Philodendrum and Alo<;mi<i
in Ecuador (vide Lagerheini in Nuova Notarisia, 1892, pp. 120124). Trichtiphiln*
Weber is a genus of Algae parasitic on the hairs of Bradypus (the Three-toed Sloth) ;

another species has also been found on species of Nenia (Clausilia); cf. Lagerheim
in Bericht. der Deutsch. Bot. Gesellsch. 1892, Bd x, Heft 8, pp. 514 517.
1
Eichter in Flora, Ixxv, 1892.
2
J. Cornere in Nuova Notarisia, xiv, 1903, pp. 1821.

July, 1902, pp. 242243.

3 G. S. West in Journ. Bot.
56 CMorophyc&z
Order II. Ohcetophorales. Thallus filamentous, sometimes
simple, but more often branched. Cells uninucleate ;

chloroplasts parietal, generally single and with pyre-

noids. Sexual reproduction either isogamous or
heterogamous. Mostly freshwater.
Order III. Ulvales. Thallus expanded, membranous, paren-
chymatous, attached when young. Cells uninu-
cleate ; chloroplasts single, parietal, with one pyre-
noid. Sexual reproduction isogamous. Mostly
marine.
Order IV. Schizogoniales. Thallus filamentous, sometimes
parenchymatous, or expanded by fusion of filaments
in one plane. Chloroplast single, central and
substellate, with one pyrenoid. Mostly subaerial.
Order V. Microsporales. Thallus filamentous, unbranched.
Cells uninucleate. with a large, parietal, reticulated
or band-like chloroplast, destitute of pyrenoids.
Exclusively freshwater.
Order VI. Cladophorales. Thallus filamentous, simple or
branched, incompletely septate. Segments large
with numerous parietal chloroplasts each with a
pyrenoid. Sexual reproduction isogamous or hetero-
gamous. Marine or freshwater.
Order VII. Siphonece. Thallus filamentous and coenocytic,
unseptate, consisting of one large branched cell
with many nuclei. Chloroplasts numerous, without
pyrenoids. Sexual reproduction heterogamous.
Mostly marine.
Order VIII. Conjugate. Thallus unicellular or filamentous.
Cells uninucleate ; chloroplasts single or several,
usually large and of some definite shape, with

pyrenoids. Sexual reproduction by isogamous

aplanogametes. Exclusively freshwater.
Order IX. Protococcoidece. Small unicellular, multicellular
or colonial Alga3. Cells uninucleate or coenocytic ;

chloroplasts very variable in form, size and disposi-

tion, with or without pyrenoids. Sexual reproduc-

tion of an isogamous or heterogamous character is

known in some. Almost exclusively freshwater.

 CEdogoniacew 57

Order I. (EDOGONIALES.
In this order the thallus consists of fixed, simple or branched
filaments. The cells possess a single nucleus and the chloroplast
a parietal, more or less cylindrical,
is
anastomosing mass of chloro-
phyll, containing one or more pyrenoids. The vegetative division,
in which a curious of new
interpolation pieces of cell-wall takes
place, is peculiar to the order.The zoogonidia are also anomalous,
being characterized by a circlet of numerous cilia round the
anterior end. In the autumn, plants of this order frequently have
their cells packed with starch. The sexual are well-dif- organs
ferentiated oogonia and antheridia, and the sexual
reproduction is
greatly specialized. There is
only one family which includes three
genera, two of which are abundantly found in the British Islands.

Family 1. CEDOGONIACE^.
This family is represented in the British Isles by numerous
species of the two widely distributed genera (Edogonium and
Bnlbochcete. The young stages of these plants possess well-de-
veloped organs of attachment, but most of the species of (Edogo-
nium float freely in the water when adult. The thallus is simple
or branched and some of the cells exhibit a peculiar transverse
striation at their upper extremities. This is particularly notice-
able in the large species of (Edogonium and is the result of inter-

calary surface growth. Beneath one of the transverse cell-walls

an annular cushion of cellulose is deposited, and after each division
a circular split is formed in the cell-wall opposite this cushion, the
two parts remaining very slightly separated by a new piece of cell-
wall derived from the cushion of cellulose. The rings or cushions
of cellulose were investigated by Hirn who found that they
1
,

consisted of a central mucilaginous mass, surrounded by a coating

of cellulose formed as an inner cell-wall layer, which becomes

intimately concrescent with the old membrane above and below

the ring. After each division another slit is formed beneath and
close to the first one, the process
being repeated until the upper
end of the frequently presents the appearance of having a
cell

number of 'caps' placed one over the other (figs. 13 B and C;

'
14 A), each cap indicating a division of the mother-cell.
'

1
Him in Acta Soc. Scient. Fennicae, xxvii, 1900.
58 Chlorophycece
Most of the cells in Bulbochcete are furnished with long
tubular bristles and the terminal cell of the filament in one or
two species of CEdogonium also ends in a long bristle (fig. 14 C).
There is one large chloroplast in each cell disposed in the
form of a cylindrical net-work, a large proportion of it forming
anastomosing cushions on the inner surface of the cell- WP 11. The
pyrenoids vary from one to several according to the species, and
sometimes the number varies in different cells of the same plant.
There is usually one nucleus with a prominent nucleolus (fig. 10 J n},
situated in a more or less central position. The nucleus occasionally
divides without a corresponding division of the cell. Growth of
the filaments takes place by the transverse division of any of the
vegetative cells.

Fig. 10. A I, (Edoponium sp., from Frizinghall, W. Yorkshire, showing stages of

one type of development from a zoogonidium in which the basal cell does not
become greatly swollen ( x 460). p, pyrenoid. J, (Edogonium sp., from Shipley
Glen, W. Yorkshire, after treatment with Acetic Acid and Hasmatoxyliu, show-
ing nuclei (n), x 460.
 CEdogoniacece 59

Asexual reproduction takes place by means of zoogonidia, which

are formed singly from ordinary vegetative cells. There is a re-
juvenescence of the entire cell-contents, a large rounded mass
being formed, which ultimately escapes. In (Edogonium this
process may take place in any of the vegetative cells of the
filament, whether terminal or not,
and it sometimes occurs in a
young plant consisting only of one
cell. The cell-wall splits into two
halves by a transverse slit near its

upper extremity and the rounded

mass of rej u venized protoplasm makes
its exit in a delicate hyaline vesicle.
This mass assumes a pyriform shape
and at the narrower end a small
colourless protuberance is formed,
round the base of which arises a
circle of numerous cilia (fig. 1 1 z}.
This striking zoogonidium, which
may or not possess a red pig-
may
ment spot, quickly swims away, the
entire process lasting only a few
minutes. On coming to rest
attacnes
,.,,,.
by anterior
itseli its
..
it
Fig. 11. The escape
from its zoogonidan-
of the zoo-
hyaline gonidiuin (z)

end, loses its cilia, and develops a

' '
8 ium - A -(Edogonium BoscU (Le
01.) Wittr., from
near Seuens, Corn-
1 his cell ultimately /
n ,.
cell-wall. forms wall. B, (E. Himii Gutw., from
a new filament by transverse cell- Churchill, Donegal, Ireland (x 460).

division (fig. 10 E I). The basal

cell may be rounded and swollen or it
may develop a hapteron or
organ of attachment (fig. D). 10 A
Wille has observed resting-spores in some species of (Edogonium '.

The sexual reproduction in this family of Alga? presents a

greater specialization of the male and female organs than is found
in any other family of the green Alga?. The oogonia may be
developed from any of the ordinary vegetative cells, and most
frequently arise from cells which exhibit intercalary surface growth
at their upper extremities. They are usually spherical or ovoidal
in form and occur singly or in series of from 2 to 10. The contents
of each oogonium become rounded off, forming a single oosphere
1
Fide Bot. Centralbl. xvi, 1883.
 CMorophyceoB
which contains much chlorophyll. The antheridia may be developed
in the same filament as the oogonia, as in the monoecious species
(fig. 12), or they may
arise separate male
in
filaments, as in the dioe-
cious species (figs. 13 and
14). The antheridia are
sometimes unicellular,
consisting of a short cell
rather narrower than the
ordinary vegetative cell

and containing less chlo-

rophyll. More frequently,
however, they consist of
more than one cell, and
occasionally of a dozen or
more, the contents of each
antheridial cell dividing
into two masses each of
which becomes an anther-
ozoid. Rarely only one
antherozoid is produced
in an antheridial cell. The
antherozoids are similar
in form to the zoogonidia
and are ciliated in the
same way, but they are
smaller and contain less

chlorophyll.
Dioecious species in
Fig. 12. Monoecious species of (Edogonium. , . , , , ,

A, a form of (E. obsoletum Wittr., froin near

which the male filaments
Goring, Oxfordshire. B, (E. zig-zag Cleve var. 3 e and
are l ar pr but little
robustum West & G. S. West, from Harefield,
Middlesex. C, (E. Itzigsolmii De Bary var. minor
.

interior
.

m
.

Size to the
West, from the Orkney Is. D, (E. Ahhtrandii fema } e filaments are said
Wittr., from Pilmoor, N. Yorkshire (x4bO).
oo, oogoniurn; a, antheridium.
to be dlOBClOUS macran-
drous (fig. 13).

There is, however, another type of dioecious species in which the

male plants are very small and are attached to the female plants ;

these are said to be dioecious nannandrous (fig. 14). This type

requires a further description. Certain short cells are produced in

 CEdogoniacece 01

the female filaments either singly or in chains, each cell being

larger than the antheridial cells of the monoecious or dioecious
macrandrous and known as an androsporangium. The
species,
androsporangiuni usually produced in the neighbourhood of an
is

oogonium and becomes the mother-cell of a motile ciliated spore

known as an androspore,
intermedia te in size between
an antherozoid and a zoogo-
nidium. Each androspore
swims about for a time and
then attaches itself to the
female plant, either actually
on the oogonium or on some
adjacent cell. It then sur-
rounds itself with a cell-wall
and grows into a very small
male plant known as a 'dwarf-
'

male or a nannandrium.
The dwarf-male usually con-
sists of a basal vegetative
cell which supports one or

more antheridial cells, but

occasionally it is reduced to
one antheridial cell only.
Two aritherozoids arise in
each antheridial cell as in
the ordinary monoecious and
dioecious species, and they
are set free by the splitting
Fig. 13. Dioecious macrandrous species of
off of a there be only
cap if
(Edogonium. A, male plant of (E. rufescens
one antheridial cell, or by Wittr., from Scilly Is. B, female plant of
same. C. female plant of (E. lautumniarum
the general dismemberment Wittr., from Welsh Harp, Middlesex. D, male
of the antheridium if there plant of same (
x 460). oo, oogonium; a, an-
theridium.
are several antheridial cells.

When the oosphere is

ready for fertilization a hyaline receptive
spot appears in it at a point opposite that part of the wall of the

oogonium which will open. The oogonium opens in many ways

but the method of opening constant for any one species. Some-
is

times a circular crack is formed, which may be median, superior,

or inferior ; sometimes a pore arises either in a superior or inferior

62 Chlorophycece

Fig.
' 14. Dicecious nannandrous species of (Edorjonium. A, a form of (E. undula-
tnm (Breb.) A. Br., from Pilmoor, N. Yorkshire. B, (E. cyathigerum Wittr.,
from Eawcliffe Common, W. Yorkshire. C, (E. ciliatum (Hass.) Pringsh.,
from near Senens, Cornwall x 460). oo, oogonium; n, naunaudrium or
(

dwarf-male; a, antheridiurn.
 (Edogoniacetr 63

position; and at other times there is a distinct

apical lid to the
oogonium. An antherozoid finds its way through the opening into
the oogonium, frequently having to accommodate itself to a
passage
much narrower than itself, and unites with the oosphere at the
region of the receptive spot. After the fusion of the nuclei of the
antherozoid and the oosphere the latter becomes the fertilized
ovum or oospore, and it
immediately surrounds itself with a cell-
wall. The oospore then rests for a longer or shorter
period, its
chlorophyll disappears, its cell-wall increases in thickness, and its
protoplasm becomes tinted with a red or brown pigment and filled
with oil. On the decay of the walls of the oogonium the
oospore
is liberatedand on germination its outer wall bursts and the
contents, surrounded by a delicate membrane, are set free. With
few exceptions a new plant is not immediately formed from the
oospore, but the free cell-contents usually divide into four cells,
each of which forms a rounded ciliated zoospore. The
zoospores
represent a rudimentary sporophyte generation, and after swarming
for a while they come to rest and form new filaments. Sometimes
the filaments formed from the zoospores are asexual and
they give
rise to several other asexual generations before
forming a sexual
plant. If the zoospores become fixed at once to some substratum,
they form a hemispherical or spheroidal cell from a circular opening
in which the new filament arises 1 . If they do not become fixed
before germination haptera are usually
developed (fig. 10 I). A
The principal investigators of this family of Algae have been
Pringsheim and Wittrock, and quite recently it has been splendid Iv
monographed by Him-.
Genus CEdogonium Link, 1820. The plants of this genus are
simple filaments with cylindrical cells usually slightly swollen at
their upper extremities. The apical cell is
generally terminated
by an acutely conical cap or more rarely by an elongated bristle.
The strong cell-walls and the swollen upper extremities of the
cells,some of which possess the peculiar transverse striation, are
characters which readily distinguish even sterile species of this
genus from all other filamentous green Alga?. The adult plants
usually occur floating in masses or they may remain attached to
various water plants, and as the mucous covering on the exterior
of the filaments is very slightly developed, they not
only serve .as
1
Scberffel in Ber. Deutsch. Bot. Ges. xix, 1901.
-
Hirn in Acta Soc. Scient. Fennicse, xxvii, 1900.
 Chlorophycece
hosts for various epiphytes, but they do not feel so slimy as most
filamentous green Algae.

There are about 80 British species of this genus, exhibiting great variation
in size and in the relative proportions of the cells. They are exceedingly
abundant, particularly in quiet waters, and with one or two exceptions the
species can only be accurately identified from fructiferous specimens. They
are frequently found in the fructiferous condition in suitable localities, such
as small ponds and ditches, and more commonly in the south of England and
south-west of Ireland than in other parts of the British Islands. The smallest
British species is (E. tapeinosporum Wittr. (diam. of vegetative cells 2'7 5/*),
and the largest (E. giganteuin Kiitz. (diam. of vegetative cells 30
is 50 /*).
(E. iindulatum (Breb.) A. Br. (fig. 14 A) possesses
very characteristic undulate

Fig. 15. A, Bulbochate subintermedia Elfv., from near Senens, Cornwall. B, B.

Nordstedtii Wittr., from near Glendoan, Donegal, Ireland. C, B. nana
Wittr., from Goring, Oxfordshire ( x 495). a, antheridium; and, androspo-
rangium ; n, nannandrium; oo, oogonium.
 (Edogoniaeeae

vegetative cells, OE. punctato-striatum De Bary has tlje entire filaments fur-
nished with spirally arranged granules, and (E. iii-rnn/iin-uni De Bary possee>
a remarkable terminal oogonium. The oospores are either globose, elliptic l,-i]

or ovoidal, and the cell-wall may be smooth, ridged, spiny, punctate, scrolm-u-
late or reticulate. Sometimes the oogonia are plicated as in (E. platygynum
Wittr., or they may possess a transversely disposed ring of conical projections
as in (E. ItzigsohniiDe Bary (fig. 12 C). In some species the supporting cell
of the oogonium is much swollen, as in (E. Borisianum (Le Cl.) Wittr. and
(E. lautumiiinim Wittr. (fig. 13 C and D).

Rather less than half the known species are dioecious nannan-
drous, and most of the remainder are monoecious.
Genus Bulbochsete Ag., 1817. The plants of this genus are
branched and every branch usually terminates in a long hollow
bristle with a swollen base. The vegetative cells widen upwards,
most of them carrying a laterally placed bristle, and they do not
reach the same length as those of (Edogonium. The
relative

oogonia are generally terminal on short lateral branches and, with ;

few exceptions, the supporting cell of the oogonium is divided by

a transverse septum, the position of which is fairly constant for

any one species. In the dioecious nannandrous species the andro-

sporangia are commonly situated on the apices of the oogonia.
The plants occur as branched tufts, more often fixed than in the

preceding genus, and they possess a quantity of enveloping mucus,

affording a home for numerous Diatoms and often Desmids.

There are about 14 British species, of which B. nana Wittr. (diam. of

vegetative cells 1015 ^; fig. 15 C) is the smallest and B. gigantea Priugsh.
(diam. of vegetative cells 24 32 /*) the largest. No doubt many more species
will be found if searched for, but in the greater part of the British Islands
fructiferous specimens are relatively scarce. There is great variability in the
form and size of the vegetative cells in the different species, and also in the

comparative size and length of the bristles. The genus is not so abundant as
(ffdagonium, and all the species prefer very still waters.

Most of the species of genus are dioecious nannandrous.

this
Few species are monoecious, and dioecious macrandrous species are
as yet unknown.

\V. A.
66 Chlorophycece

Order II. CH^TOPHORALES.

In this order of green Algse the thallus is filamentous, sometimes
simple, but more frequently branched. The branches are generally
attenuated and often piliferous. The cells possess one nucleus,
and in the families of the order, except the Trentepohliaceae,
all

there isa single parietal chloroplast with one or more pyrenoids.

Asexual reproduction takes place often by resting-spores, which
may be either aplanospores or akinetes, and
commonly by zoogonidia
with two or four cilia. Sexual reproduction is brought about
by
isogamous planogametes with two cilia, or by well -differentiated
heterogamous gametes.
This order has also received the name of the " Ulotrichales,"
but I prefer to accept Wille's name of the " Chsetophorales " as
five out of the seven families include branched
Algae.

Family 1. Coleochcetacece. Flat expansions or pulvinate branched

masses, epiphytic on the stems and leaves of submerged plants. Sexual
reproduction heterogamous plants mono3cious or dioecious oogonia
; ;

with a trichogyne and one non-motile oosphere fertilization within;

the oogonium and resulting in the formation of a cortical layer on the

outer surface of the oogonium. Some of the cells of the thallus are
furnished with fine bristles with basal sheaths.

Family 2. Herposteiraceoe. Filaments branched, creeping, epi-

phytic on submerged plants. Sexual reproduction heterogamous ;

plants monoecious oospheres motile, fertilization taking place outside

;

the oogonium. Cells with one or several long bristles, sometimes

swollen at the base.

Family 3. Ulotnchaceoe, Filaments simple. Chloroplast single,

parietal, with one or many pyrenoids. Sexual reproduction isogamous.

Family 4. Ci/lindrocapsacece. Filaments simple cells with thick ;

lamellose coats, usually arranged in a single series within a lamellose

gelatinous sheath. Sexual reproduction heterogamous plants monoe- ;

cious ; oogonia with one non-motile oosphere ;

fertilization within the

oogonium.
Family 5. Chcetophoracece. Filaments branched branches attenu- ;

ated into multicellular hair-like prolongations. Chloroplast .single,

parietal, with one or many pyrenoids. All the cells except those of the
rhizoids and hairs are capable of producing zoogonidia or
gametes.
Sexual reproduction isogamous.

Family 6. Microthamniaceas. Filaments branched; branches

scarcely attenuated, not piliferous. Chloroplast single, parietal, with
 Coleoehcetaeece 07

or without a single pyrenoid. Zoogonidia and gametes produced in

special gonidangia. Sexual reproduction isogamous.
Family 7. Trentepohliacece. Thallus branched, terrestrial or ar-
boreal. Chloroplasts several, parietal, without pyrenoids. Zoogonidia
and gametes produced in special gouidangia. Sexual reproduction
isogamous.

Family 1. COLEOCH^ETACE.Sl.
The plants included in this small family have reached a higher
stage of development than any other of the green Alga?, and have
undoubtedly arisen from the Chsetophoracea? by further speciali-
zation. They form small discs or cushion-like masses, which are

enveloped in mucilage and are attached to the stems and leaves of

larger water-plants. In the commoner forms the thallus is more
or less circular in outline and disc-like in form, consisting of a

Fig. 16. Coleochcste scutata Breb. (

x 100), from Welsh Harp, Middlesex.

single layer of cells in one plane, which either form a compel

parenchymatous layer, or are arranged in the form of branched
filaments radiating from a central point. In other species the
ramification is not confined to one plane, but numerous
ascending
branches are given off, the whole thallus sometimes
having the
appearance of a hemispherical cushion. The peripheral cells of
the disc or the terminal cells of the branches are meristematic and
the thallus grows by the formation of new radial and
tangential
cell-walls. The branching is in some species dichotomous, but in
others it closely resembles that of
Chcetophora or Myxonema.
52
08 ChlorophycecB
Some of the cells are furnished with a colourless bristle which is

fixed at its base into a narrow sheath of considerable length.

Asexual reproduction takes place by means of large ovoidal
zoogonidia (fig. 17 D), furnished with two long cilia and produced
singly from the cells of the thallus, more particularly from the
terminal cells of the branches. The zoogonidium escapes from
the zoogonidangium either by a round orifice on the upper surface
or by the dissolution of the extremity of the terminal cell.
Sexual reproduction is brought about by the fertilization of an
oosphere by an antherozoid. The sexual organs are oogonia and

Fig. 17. Coleoclw>te pulvinata A. Br. A and B, from near Glenties, Donegal, Ireland;
A, portion of thallus with sexual organs ( x 460) o, oogoniuin
; ;t, trichogyne ;

B, ripe spermocarp emitting the cells formed by the division

'
'
, autheridia.
of the oospore each of these becomes a zoospore ( x 4GO). C, zoospore (after
;

Chodat). D, zoogonidium (after Pringsheim).

antheridia. The oogonium is developed by a swelling of the

terminal cell of a branch and it possesses on its upper surface a
narrow trichogyne. An oosphere containing chlorophyll is pro-
duced within the oogonium, and just previous to fertilization the
of
trichogyne opens at the apex and exudes a colourless drop
mucilage. The antheridia are flask-shaped cells which are developed
from cells in the neighbourhood of the oogonium, or in dioecious
species from cells of another thallus. Only one antherozoid is
 ColeochcetacecB Of)

produced in an antheridium and it can only be distinguished from

a zoogonidium by its smaller size. After fertilization the oospore
surrounds itself with a cellulose wall and grows considerably in
size. At the same time the oogonium becomes closely covered
with a layer of cortical cells, produced by the proliferation of the
supporting-cell and by the close application of the terminal cells of
other branches. The fertilization and the formation of this cortical
layer are said by Pringsheim to take place from May to July. The
whole structure produced after fertilization, and which presents
the appearance of 'a sphere supported on one or many filaments,
has been termed a "spermocarp." The cortical cells often become
dark brown or red in colour and lose their chlorophyllaceous con-
tents. Usually this structure remains dormant through the winter,
the maturation of the oospore taking place slowly. On germination
the oospore divides into a number of cells and the cortical layer
splits irregularly into two halves. The escaping spores (fig. 17 B),
which become more or less irregular in outline, do not give rise
directly to a new thallus, but each one becomes a zoospore with
two cilia. The zoospore (fig. 17 C) gives rise to several rudi-
mentary asexual generations which are propagated by zoospores,
and finally to a sexual individual.
Genus Coleochaete Breb., 1844. The thallus is filamentous,
branched, erect or usually forming a fiat pseudo-
creeping,
parenchymatous plate with peripheral growth. The bristles, which
are sparsely scattered over the upper surface of the thallus, are
not always clearly visible, and they are characterized by the well-
marked sheathing base. The plants are all epiphytes with a
marked dorsiventral development, but there are no special organs
of attachment. Each vegetative cell possesses a large nucleus and a

single parietal chloroplast of irregular form, which contains one or

two large pyrenoids. The plants occur attached to the submerged

portions of various aquatic and marsh plants from which they niv
not easily removed.
C. scutata Breb. and Pringsh. are the most abundant species in
C. soluta
Britain. The former possesses a compact, flat, parenchymatoua thallus
(fig. 16)
and the latter a flat thallus composed of dichotomously branched
filaments radiating in one plane from one or more central cells. The diameter
of the thallus in each case scarcely exceeds 700 800 p and the cells average
abovit 10 23 /x in diameter. ('. orbicuf art's Pringsh. possesses a flat, expanded,
circular thallus which reaches a diameter of 4 mm., in which the filaments
are very closely packed and the cells are rather small. C. pulvinnta A. Br.
70 Chlorophycew
forms hemispherical cushions commonly 2 4 mm. in diameter, but occasion-
ally greatly exceeding these dimensions.
The filaments are erect and radiating
and the cells are 1 3 times longer than their diameter, which is from 20 50 p.
(fig. 17).
This species is more frequently observed with sexual organs than
any of the others. C. irregularis Pringsh. possesses a more or less parenchy-
rnatous thallus in which the branching is very irregular. All the species are
readily eaten by pond-snails of the genera Limnwi and Planorbis.

Family 2. HERPOSTEIRACE^E.
Thisa small family including only the genus Herposteiron.
is

The plants are epiphytic on larger Algse and on other water-plants,

and occur as short irregular filaments which are little branched.
Most of the cells of the filament possess one or more bristle-like
seta? or hairs, cut off from the cell which bears them by a basal

Fig. 18. Herposteiron confervicola Nag. (=Aphanochcete repens A. Br.). oo,

oogonium ;os, oospbere;a, antheridium an, spermatozoid.
; (After Huber.)

septum. Chodat has found that in cultures the setae are sometimes
replaced by branches, showing the relationship
between this genus
and the Chsetophoracese.
Asexual reproduction takes place by zoogonidia, one to four
being produced from the mother-cell, the wall of which ruptures
and sets them free. They vary much in size, possess four cilia,
and usually a red pigment-spot. On coming to rest they generally
develop unilaterally into a new plant. Sometimes aplanospores
are formed (fig. 19 Ca).
The sexual reproduction of Herposteiron is of special interest.
The oogonia are differentiated from certain of the central cells of
 Herposteiracew 71

the thallus which are devoid of bristles. These cells grow in size,
assume a globular form, and become filled with starchy and oily
material. One oosphere is produced, which is motile, having i'mn
cilia, and is expelled from the oogonium by the rupture of the

upper portion of the wall. The antheridia are small cells usually
developed at the ends of the filaments and branches they are;

frequently colourless and are considerably smaller than the ordinary

vegetative cells. One or two antherozoids are produced in an

Fig. 19. A, Herposteiron pilosissima (Schmiclle) nob., from Wimpole Park, Cam-
bridgeshire. B D, H. confervicolfi Nag. B arid C, from Bradford, W. Yorks.
; ;

D, from Eichmoud Park, Surrey ( x 450). a, aplanospore.

antheridium. These are pear-shaped bodies with four cilia and

two pulsating vacuoles, and are much smaller than the zoogonidia.
They escape into a hyaline vesicle which soon becomes diffluent
and sets them free. The antherozoids move about very rapidly,
but the movements of the oosphere are very feeble. The Herpo-
steiracea? is the only family of the Chretophorales in which the
fertilization of the oosphere takes place outside the oogonium.
Little is known the development of the oospore.
concerning
Genus Herposteiron Nag., 1849. [Aphanochcete A. Br., 1851 ;

Berth., 1878 Huber, 1892.] The thallus is filamentous, creeping

;
72 Chlorophycece

and branched, the terminal cells of the branches being rather

smaller than the more central cells of the thallus. One or more
erect bristles are attached to the dorsal surface of some or all of

the cells. These bristles are single, attenuated, and very elongated
cells, which have lost their protoplasmic
contents and which never

possess chloroplasts. They are somewhat fragile and ar^, easily

broken off near the base. The sexual organs, which are of very
rare occurrence, have been described by Huber
1
.

The plants occur as epiphytes on species of (Edogonium,

Cladophora, Rltizoclonium, and Mougeotia, and

also on the leaves

of Lemna, Elodea, etc. Sometimes well-developed specimens are

much branched, the procumbent branches of the Alga often
of the plant to which it is
following the contours of the cells
attached, and in consequence exhibiting a marked reticular
structure 2 (fig.
19 B). The cells contain a single parietal chloro-

plast with one (or

more ?) pyrenoids.
There has been much confusion with regard to the two names

Aphanochwte and Herposteiron. The arrangement proposed by

subsequently adopted by De Toni, Wille
and and others
Hansgirg,
of two distinct genera is quite untenable. There
(myself included),
can be no doubt in the mind of anyone who has studied these
and Aphano-
plants carefully that Herposteiron confervicola Nag.
chcete repens A. Br. are descriptions of the same plant, and this is

amply confirmed by the authentic drawings by Nageli published

admit the identity of
by Huber. Both Huber and Klebahn
reject Nageli's name
and Aphanochcete, but on the
Herposteiron
of the incompleteness of the description. There is, however,
ground
far more reason for neglecting Braun's name on the ground of

inaccuracy.

H. confervicola Nag. (
= Aphanochcete repens A. Br.) is a species with
oblong-ellipsoidal cells, each bearing a single bristle which is little swollen at
the base and which is attached towards one end of the cell. It is not an un-
common species and is somewhat variable,
two bristles being frequently
attached to some of the cells of the thallus. (Figs. 18 and 19 B D.) Another
nob. ( = ApkanoeJuete pilosissima Schmidle),
species, H. pilosissima (Schmidle)
is more abundant in some parts of the British Islands and is most probably

identical with H. Hansg. The cells are more ellipsoidal and possess
polychcete,
from one to four bristles, each bristle having a swollen base (fig. 19 A).

]
Huber in Bull, de la Soc. bot. de France, xli, 1892.
a
G. S. West in Journ. Bot. Febr. 1899, p. 57.
 Ulotrichacece ?:>

Family 3. ULOTRICHACE^E.
This family includes a few genera which are readily distin-
guished from other plants of the Chsetophorales by their un-
branched habit and by the structure of their cells. The thai! us
is a simple filament, consisting of cylindrical or doliform cells, as
in Ulothrix, or of rounded cells arranged in a single series and
enveloped in a thick mucous coat, as in Hormospora and Radio-
filum. The cell-wall is always hyaline and colourless, but varies
much in thickness. It is sometimes delicate, sometimes thick and
lamellose, and sometimes the outer layers are diffluent. There is

a single, parietal, plate-like chloroplast in each cell, with an entire

or variously lobed margin, and containing one or many pyrenoids.
A single nucleus is
present in the cytoplasm.
Asexual reproduction takes place in several ways. Sometimes
aplanospores are produced (fig. 20 D a fig. 21 F a), or numbers
;

of akinetes are formed by the enlargement of certain cells and the

1

gelatinization of the outer portions of their original cell-walls

(fig.
21 E and I); these may be resting-spores (hypnospores) or

they may germinate directly. Sometimes the thallus is multiplied

by a general dismemberment of the filament into single cells or
groups of cells, each cell or group developing into a new filament.
Zoogonidia of two kinds are produced, often from different cells of
the same filament; small microzoogonidia with two cilia and
larger macrozoogonidia with four cilia. The microzoogonidia are
produced from certain of the vegetative cells which have become
microzoogonidangia and in the larger species of Ulothrix, such as
U. zonata, 16 or 32 are produced from each gonidangium, but in
U. subtilis only 2 or 4 are produced. Similarly 2, 4, or 8 macro-
zoogonidia are usually produced from a macrozoogonidangium, but
in U. subtilis only one arises. It is occasionally observed that the
entire contents of the cell are not used up in the formation of the
macrozoogonidia {vide fig.
21 G). The zoogonidia germinate di-
rectly on coming sometimes even within the mother-cell,
to rest,
and the plants which by the germination of the macro-
arise

zoogonidia are larger than those which arise from the micro-
zoogonidia. This accounts for the variability in size of tin-
filaments which is so often observed in a collection of any one

1
This was first shown by Wille in Bot. Centralbl. xi, 1882, p. 113.
74 Chlorophycece

species of Ulotlirix. Occasionally the zoogonidia do not escape,

but lose their cilia, become invested with a cell-wall, and form
what is termed a "palmelloid condition" (fig. 20 F). The produc-
tion of zoogonidia usually commences near the apex of a filament
and progresses towards the base.
Sexual reproduction is by the conjugation of isogamous gametes,
which are indistinguishable from the microzoogonidia. The gametes
are biciliated and usually escape from the gametangia in the

morning, conjugating in pairs with considerable rapidity. The

Fig. 20. A and B, Ulothrix zonata (Web. et Mohr.) Kiitz., from near Means Abbey,
E. Yorkshire ( x 500). C F, U. subtilis Kiitz., from near Mullion, Cornwall
(x-500); F sbows the "palmelloid condition"; a, aplanospore za, rnacro-
;

zoogomdium zi, microzoogonidium.

;

resulting zygospore invests itself with a firm cell-wall and germi-

nates after a more or less extended period of repose. On germina-
tion the contents break up into many zoospores each of which
forms a new filament. The gametes frequently germinate directly
without conjugation.
The movements and gametes are fre-
of the microzoogonidia

quently very strange, one cilium being kept more or less rigid and
its extreme apex used as a pivot, while the other cilium exhibits

violent movements causing a rapid lateral oscillation of the

body.
 Ulotrichacece 75

The British genera are best arranged as follows :

* Filaments thread-like, cells cylindrical with truncate

apices.
t Filaments long and flexuose, attenuated towards
base Ulothrix.
ft Filaments short, attenuated at both base and apex Uronema.
ftt Filaments of variable length; transverse walls
very thick; cells in pairs Binuclearia.
** Filaments fragile, often moniliforin, cells with rounded
apices.
t Cells more or less cylindrical ; plants with a re-
semblance to a fragmented with no
Ulothrix;
prominent mucous coat Stichococcus.
ft Cells cylindrical with hemispherical ends, or sub-
globose, often remote with a prominent mucous
;

envelope.
Cells cylindrical.

| Cells equidistant, often in close contact.

Cells large, short Hormospora.
Cells minute, more elongate CloeotiJu.

H Cells in pairs Geminella.

( 'ells rounded Radio/Hum.
The three genera Hormospora, Glceotila and Geminella are scarcely to be
distinguished from each other. Perhaps it would be better to unite them
under the name Geminella.

Genus Ulothrix Klitz., 1833. [Hormiscia in the sense used

by Rabenhorst (1868), Hansgirg, and De Toni.] In this genus
the filaments are simple, not attenuated at the apex, but fre-
quently fixed at the base by a unicellular, simple or ramified
'
rhizoid.' The cells are commonly cylindrical or sometimes swollen,
and in the larger species the cell-wall is thick and evidently lamel-
lose. The chloroplast is parietal with one or many pyrenoids, and
varies much in its relative size.
The genus Ulothrix was established by Ktitzing 1 for the species
U. two years
zoimta before Fries' description of Hormiscia-.

Areschoug's enlargement of the genus Hormiscia was based upon

3

erroneous conceptions, as he included in it species having no

affinity with each other.
The original Hormiscia of Fries only
included two Algae previously known as "Conferva penicilliformis
Roth" and "Conferva Wormskioldii Flor. Dan." These Alga? are
commonly placed under the genus Urospora of Areschoug, but the
1
Kiitzing in Flora, 1833, xvi, p. 517.
2
Fries in Flora Scand. 1835, p. 327.
3 no.
Areschoug in Acta Beg. Soc. Sci. Upsala, ser. in, vol. vi, 2, p. 12.
76 Ctilorophycece

latter genus should undoubtedly be placed as a synonym of Hor-

miscia Fries.
The best known species of the genus is U. zonata (Web. et Mohr.) Kiitz.,
(tig.
20 A and B), which is widely distributed all over the British Islands,

occurring as bright green masses in streams, rivers, etc., more especially

in
the early spring. The cells vary from 15 70 /x in diameter and the cell-walls
are very thick and lamellose. An abundant British species is U. subt Hi's Kiitz.

Fig. 21. A F, Ulothri.r a-qualls Kiitz.; A D, from Putney Heath, Surrey; E and
F, from Mitcham Common, Surrey; A, filament showing escape of micro-
zoogonidia B D, germinating macrozoogonidia, C shows the same plants as
;

B 48 hours afterwards, D is much further advanced; E, portion of filament of

akinetes F shows two aplanospores which have taken exactly 14 days to
;

develop from ordinary vegetative cells. G, U. vyi///.s Kiitz. var. catceniformis

(Kiitz.) Rabenh., from near Bradford,
W. Yorkshire, showing escape of macro-
zoogonidia. H, U. moniliformis Kiitz., from Wimbledon Common, Surrey ;

I, the same with akinetes. (All x 500. a, aplauospore

)
ak, akinete ; za, macro-
;

zoogonidiura ; zi, microzoogonidium.

(fig.
20 C F), the cells of which are as long as broad and from 4 8 fj.
in

diameter. A
variety of this species var. variabilis (Kiitz.)
Kirch n. is

probably the most abundant member of the genus, being generally

distributed
in the stagnant waters of ponds, ditches, troughs, rain-tubs, etc. It is a little
thicker than U. subtilis and the cells are lj 2j times longer than their
diameter. U. cequalis Kiitz. (fig. 21 A F) and U. moniliformis Kiitz. (fig. 21
H are other well-known species.
I)
Another Alga Sclnzomeris Leibldnii Kiitz. which I have only once seen,
from Stone Ghyll, Dodd Fell, N. Yorkshire, should perhaps be included here.
I am doubtful as to the exact determination of the Yorkshire specimens, but

they reminded one very much of a large Ulothrix zonata, attenuated both at
the apex and the base, the latter being fixed to rocks and stoues in the spray
of a waterfall. Longitudinal division of the cells had occurred at intervals, so
 Ulotrichacece I I

that the filaments often consisted of a double row of cells. Wolle seems
to have observed the same plant from several
parts of the United States
(cf. Freshw. Alg. of U. S. t. cxxv).

Genus Hormospora Breb., 1840. The filaments are simple

and free-floating, rarely fixed by a mucous disc, and they consist of
a single series of cells embedded
in a thick, cylindrical, mucous in-
vestment. This outer gelatinous
coat varies in its relative size,
8
and is
always hyaline and homo- 9
geneous. The cells are
commonly B
oblong-cylindrical with broadly-
rounded extremities, and they
contain a single parietal chloro-
plast usually disposed as an
equatorial band. One pyrenoid
is
generally present in each chlo-
roplast, although rarely pyrenoids
are quite absent. The genus is

very closely allied to Ulothrix,

but distinguished by its thick
is

gelatinous coat and by the con-

Fig. 22. A, Hormospora mutaliUix
stant separation of the cells after Breb., from near Mullion, Cornwall.
division. Cienkowski and others B, H. ordinata West & G. S. West,
from Cam Fell, W. Yorkshire. C E,
have regarded the genus as a mere Glceotiln protofieiiita Kutz., from Pil-

state or condition of Ulothrix, but moor, N. Yorkshire. ( x 440.)

I think that open to much doubt. The cell-wall is extremely thin

is

and delicate, and the formation of zoogonidia has not been observed.
The most frequent British species is H. mutabilis Breb. (fig. 22 A), which
occurs principally in bogs, especially amongst Sphagnum, and in such
localities species of Ulothrix do not usually exist. The cells are 16 19 p
in diameter and \\
If times longer than broad. H. ordinata West &
G. S. West 22 B) is a smaller and much rarer species with
(fig. cells 5 -8 p.
in

diameter. H. plena Breb. is the only other British species.

Genus Glceotila Kutz., 1843. This is a genus of small Alga 1

intermediate in character between Hormospora and Ulothrix.

The cells are very small, oblong or elliptical and more or less
moniliform, but they are not so completely separated as those of
Hormospora they are arranged in a single series in a delicate
;

mucous envelope, and each one contains a parietal chloroplast

of small size which I have not
is
disposed as in Hormospjra.
 78 Chlorophycece
observed any pyrenoids in the chloroplasts of this genus. Borzi 1
has recently described the formation of zoogonidia.
The cells of G. protogenita Kiitz. (fig. 22 C E), which is the typical species
of the genus, only reach a diameter of 3 4'5 p.. The plants are very rare and
occur in bogs or boggy pools.
Genus Geminella Turp., 1828 s Lagerh., 1883 3 ;
.
[1 Planctonema
Schmidle, 1903.] This genus is scarcely to be distinguished from
Hormospora Breb. except for the arrangement of the cells in pairs.
The cells are fairly remote, are enveloped in a thick mucous coat,
and after division the daughter-cells separate only very slightly.
The chloroplast is exactly as in Hormospora. The cells of some
of the filaments occasionally develop thick brown cell-walls arid
become resting akinetes.
G. interrupta Turpin (fig. 23 A
the only known species and it is
C) is

rarely found in the British Isles. are 6'5 7 The

in diameter.
cells
It /j.

would perhaps be more correct to unite the genera Geminella and Hormospora,
the former having priority.
It is impossible to find any

generic characters sufficient to

separate Planctonema Schmidle
from Geminella or Grlveotila.
Genus Radiofilum Schmidle,
1894. The filaments are simple,
sometimes short and fragile, some-
times long and flexuose, and they
H are enclosed in a considerable
mucous sheath, which exhibits
a more or less distinct radiating
fibrillar structure. The cells are
globose, ellipsoid, or sublenti-
cular, free and distant or joined

by a narrow hyaline bridge, always

I forming moniliform filaments
after the manner of those of the
Fig. 23. A C, Geminella interrupta
Turp. A and B, from near the Lizard,
;
Nostocacese. In each cell there
Cornwall ( x 440) C, two resting aki-
;
one chloroplast containing a
is
netes from Glen Tummel, Perthshire,
Scotland ( x 350). D, RadiolUnm flaves- single pyrenoid. In one species
G. S. West, from Wicken Fen,
composed of two
cciis the cell-wall is
Cambridgeshire (x440).
'
1
Borzi, Studi Algologici II.'

Turpin in Mdm. du Mus. d'hist. nat. 1828, torn, xvi, p. 329, t. 13, f. 24.
3
Lagerh. in Ofvers. af K. Vet.-Akad. Forh. 1883, no. 2.
 Ulotrichacece

equal halves, but in others it is not. The cells multiply by division

which is preceded by a division of the chloroplasts and pyrenoids.
The type species of the genus, R. conjunctioum Schmidle 1 has not been ,

observed from Britain. K. flavescens G. S. West (fig. 23 D) is a larger

species, with much longer flexuose filaments and
broad elliptical cells the ;

diameter of the cells is 7'5 10'5 /* and the chromatophores are of a yellowish-
green colour.
Genus Stichococcus Nag., 1849. [Hormococcus Chodat, 1902.]
The filaments are entirely or very largely aerial, and are composed
of cylindrical cells. They readily become disarticulated into frag-
ments composed of a few cells, the extremities of the terminal
cells being broadly rounded. There
is one parietal chloroplast in

each cell, plate-like or or more

less irregular in form, and occupying
as a rule only a portion of the cell-wall. small pyrenoid is A
present in each. In some of the aquatic forms of Stichococcus

flaccidus I have observed a fragmenta-

tion of the chloroplast (cf. fig. 24 B).
The plants are propagated by
division of the cells and fragmen-
tation of the filaments, by akinetes,
and by zoogonidia. The latter are bi-
ciliated, have no pigment spot, and
arise singly from cells which are under-

going rapid division.

This genus much resembles Ulothrix,
but its adaptation to an aerial existence
has caused a multiplication by dis-
articulation of the filaments and a
reduction almost to a unicellular con-
dition. This disarticulation often takes
place first on one side and then on the
other, giving a zig-zag appearance to
the disarticulated filaments. Klebs has 24. A, Stichococcus
bacillaris Nag., from Saltaire,
shown that it is facilitated either by W. Yorkshire. B, S. flaccidus
too much or too little nourishment. (Kiitz.) Gay, from Barnes Com-
2
mon, Surrey. C, S. dissectus
The genus was well studied by Gay .
Gay, from damp walls, London.
There appears to be no justification D, S. rtiriabilis West & G. S.
West, from Bradford, W. York-
whatever for Chodat's name "Hormo- shire ( x440).
coccus."
1
Schmidle in Flora, 1894, Heft 1, p. 47, t. vii, f. 4, 5.
2
Gay, 'Recherches sur le devel. et les classif. de quelques Algues Vertes,'
Paris, 1891.
80 Chlorophycea'
8. baeillaris Nag. (fig. 24 A) is an abundant species on damp earth, walls,
palings, etc. ;
diam. of cells '1 S.flaccidus (Kiitz.) Gay (fig. 24 B) is
3'8 p.
a larger species frequent on wet stones and in rain-pools; diam. of cells
710-5 8. dissectus Gay (fig. 24 C) is a closely allied species to 8. Jlac<:iihis,
fj..

or perhaps only a form of it. S. variabilis West & G. S. West 24 (fig. D)

forms a thin green stratum on wet stones in the neighbourhood of waterfalls ;

the cells are very irregular in outward form and the chloroplast is often devoid
of a pyrenoid or may even possess two diameter of cells 3; 6 p.

Genus Uronema Lagerh., 1887 The filaments are simple,

1
.

relatively short, and destitute of a mucous coat they consist of ;

cylindrical cells, the apical cell being acuminate and the basal cell
attenuate. The plants are fixed by a disc secreted by the basal
cell. The chloroplast occupies a con-
siderable area of the cell-wall and is

parietal, containing two pyrenoids. The

cell-wall is firm and thin.
The zoogonidia are produced singly
or in pairs from each cell; they possess
four cilia and a subapical pigment-spot.
Sometimes the zoogonidia are arrested
in their escape, the cilia are not de-

veloped, and an aplanospore is produced

by the acquirement of a strong cell-
wall.

The only species is U. confer vicoluin Lagerh.

with filaments 4 6 p. in diameter and cells
2 3 times longer than broad. It is an ex-

ceedingly rare plant, distinguished from species

of Ulothrix by its short filaments, by the
attenuation of the apical and basal cells, by
the chloroplast, and by the firm cell- walls. I
have only met with it in abundance from the
Fig. 25. A E, Binuclearia
Wittr., from Lewis,
tiitrini/1 Orkney and Shetland Is.
Outer Hebrides ( x 440).
The genus Rhaphidonema Lagerh.
(ofwhich R. nivale is known from W. Yorks.) is a fungus.
Genus Binuclearia Wittr., 1886 2 The filaments are simple .

and attached when young by a hapteron from the basal cell. The
cells are cylindrical with firm, distinctly lamellose cell-walls, the
transverse walls being unequal, a thin one and a very thick one
alternating. The cells thus appear to be arranged in pairs. The
1
Lagerh. in Malpighia 1887, p. 518, t. xii, f. 110.
-
Wittr. in Wittr. and Nordst. Alg. Exsic. 1886, no. 715. See also Schroder in
Forschimgsberichte aus der biol. Station zu Plon, Teil vi, 1898, p. 1921.
 Cylindrocapsaeece 81

chloroplast is
single, parietal, and disposed as part of an equatorial
band. Wittrock described the presence of two granule-like bodies
of a nutritive character, which he termed "nuclei," situated one
towards each end of the cell and outside the chloroplast. I have
examined quantities of this plant and find those bodies commonly
absent.

B. tatrana Wittr. (fig. 25 A E) which has cells 6 9 \L in diameter, occurs in

mountain lakes and bogs. It resembles certain stages of species of Tribonema
(Conferva), and forms hypnospores (cfr fig. 25 E), but is more rightly placed
in the Ulotrichacese on account of its parietal chloroplast. Sometimes the
filaments become distinctly mucous.

Family 4. CYLINDROCAPSACEJE.
This family includes only a few plants belonging to the genus
Cylindrocapsa Reinsch. The thallus is filamentous and un-
branched, and resembles very much that of certain of the Ulotri-
chacese. The cells are disposed in a single series, each one being
surrounded by a lamellose, gelatinous cell- wall, and the entire
filament is enclosed in a thick lamellose sheath. The cells
resemble very much
those of the genus Hormospora in their
disposition and they may divide in the same manner as those of
Radio/Hum; they are often ovoid or subtriangular in shape and
disposed in pairs at intervals along the filament. Each cell

possesses a parietal chloroplast with a single pyrenoid, but it is

often difficult to observe the nature of this chromatophore.

Asexual reproduction occurs by zoogonidia formed singly, or
in twos orfours, from any of the cells of the filament. Each
zoogonidium is rounded or oval in form, possesses two cilia, a red
pigment-spot and two contractile vacuoles.
Sexual reproduction takes place by means of well differentiated
male and female gametes. The male organs or antheridia are the
result of the active division of certain vegetative cells, and are

disposed in one, two, or four longitudinal series within the lamel-

lose sheath. Two antherozoids are produced in each antheridial
cell, similar in form to the zoogonidia, brownish red in colour, and
with two short cilia. The oogonia are developed by an increase in
size of the ordinary vegetative cells, each oogonium being large,
ovoidal in shape, and with a thick lamellose wall. A single
oosphere is
present in each oogonium, which opens by a lateral

w. A. 6
82 Cfilorophycece

pore admit the antherozoids. On

to fertilization the oospore
does not
develops a brick-red colour and
a thick cell-wall, but it

ii

1
a
, -_ .
.^.
. ." o ?-\*

A D, Cylindrocapsa involuta Reinscb (x480). a, antheridium an,

Fig. 26. ;

antherozoid oo, oogonium.

; (After Cienkowski.)
E and F, C. conferta
West, from Bowness, Westmoreland (x520j.

fill the oogonium. The method of sexual reproduction was worked

1
out by Cienkowski .

Genus Cylindrocapsa Reinsch, 1867. The thallus consists of

unbranched filaments of cells, each cell having a thick lamellose
cell-wall. The filaments are encased in a thick lamellose sheath,
and they greatly resemble certain of the more gelatinous stages
1
Cienkowski in Bull, de 1'Acad. Imp. St Petersbourg, torn, xxii, 1876, pp. 549
555, t. ii, f. 5065.
 Chcetophoracece 83

met with in the Ulotrichacese ;

in tact, many authors place this
genus in the Ulotrichacese.
Species of this genus are rarely met with in the British Isles. C. involuta

Reinsoh (which includes C. nuda Reinsch), the cells of which are 23 30 p. in

diameter, is known from Ireland (fig. 26 A C) C. conferta West (fig. 26 E
;

and F) is known from the English Lake District, and C. geminella Wolle var.
minor Hansg. has been observed from Yorkshire and Cambridgeshire.

Family 5. CH^JTOPHORACE^J.
This family of the Choetophorales has undoubtedly arisen by a
further specialization of the Ulotrichacere. The thallus is branched
and the branches are attenuated, sometimes being produced into
long multicellular hairs. As a rule the thallus is
differentiated
into a recumbent or creeping portion, attached to a substratum by
rhizoids,and an erect, branched portion. The creeping portion
presents a more or less moniliform
or torulose appearance, is

branched, and the cells are very similar to those of the Pleurococ-
caceae. The the erect portion of the thallus are elongated,
cells of

more or less swollen, but not torulose, and the branching is most
irregular, the terminal cells of
the branches frequently forming

long hyaline hairs.

There is a single chloroplast in each consisting of a
cell,

parietal, more
or less irregular plate, containing one or more

pyrenoids. In the attenuated cells towards

the ends of the branches
the chloroplast becomes reduced, and in the long, hyaline, terminal
cells it is entirely absent.
i/

Zoogonidia may be produced from all the cells of the thallus

except those forming the rhizoids or the terminal hairs of the

branches. The number which may arise from a single cell varies
from 1 to 16, depending upon the age of the plant, the size of the

cell, and other indeterminable causes. Both macrozoogonidia and

microzoogonidia are produced, exhibiting a considerable range in
size, and they possess either two or four and a pigment spot.
cilia

They rapidly come to rest, lose their cilia, and germinate directly.
On the direct germination of a zoogonidium the cilia are lost, a
cell-wall arises, and increase in length takes place, one pole being

greatly elongated to form a hyaline, rhizoid-like projection. Septa

soon appear dividing the original long cell into several shorter
ones, and if the adult plant be a strongly branched one, the
62
84 Chlor.ophycece

branching soon becomes apparent. The zoogonidia in both Chce-

tophora and (Stigeocloninm) frequently congregate in

Myxonema
masses on becoming quiescent and almost all germinate simul-
taneously (vide fig. 28 G). Palmelloid groups sometimes arise
in Myxonema by the de-

generation of some of the

branches. Famintzin and 1

Fritsch 2 have observed these

palmelloid cells germinate

directly to form new plants,
3
and Cienkowski has seen
them give rise to microzoo-
gonidia, which latter form
the young plants.
Akinetes( which are rest-
ing-cells or hypnocysts) are
frequently produced in all
the genera of this family.
In this condition of the plant
almost all the cells of a tuft
of branches take part in

spore-formation, one rest-

ing-spore being formed in

each The original cell-
cell.

walls become hyaline or in-

distinct, causing the branch-
es to exhibit a moniliform
appearance. Each akinete
is of a red-brown colour with

a thick, asperulate cell- wall

(fig.
29 D).
The gametes are small
biciliated bodies, practically

indistinguishable from the

Fig. 27. ABndB.CAdrtopAofatnmiMata
from
(Hurts.) Hazen, Scarborough Mere, microZOOgonidia except for
the possession of only two
(nat. size). cilia, and they conjugate in

265.
Famintzin Melang. Biol. Bull. Acart. St Ptersbourg, torn, viii, 1871, p.
1
in
-
Fritsch in Beihefte zum Botauischen Centralblatt, 1903, Brt xin, Heit 4,
p. 384.
3
Cienkowski in Botan. Zeitung, 1876, xxxiv.
 Chwtophoracece 85

pairs. The resultant zygospores usually undergo a short period

of rest before germination.
Genus Chaetophora Schrank, 1789. The thallus is gelatinous,
macroscopic, of a tough consistency, and of some definite form.
The filaments radiate out from a central point, those nearest the
centre being little branched, but carrying at their apices dense

clusters of corymbiform branches of a very bright green colour.

The terminal cells of the branches are often prolonged into long
hyaline hairs. The zoogonidia possess two or four cilia, and the
hypnospores are brown, being generally developed from the terminal
cells of the branches. Many of these plants, particularly certain
1
species have the power of extracting calcium carbonate from the
,

water in which they live and so giving rise to incrustations of

considerable thickness.
The most abundant species of the genus is Oh. pisiformis (Roth) Ag., an
Alga which occurs as hemispherical, or almost spherical, dark green masses
attached to submerged stones or to the submerged parts of plants. It is also
often found attached to the shells of aquatic Gastropods. Its distinguishing
features are the absence of terminal hairs and the slightly torulose character
of cells of the branches; the cells of the primary filaments are 9 15 p in
diameter. The next most abundant species is Ch. incrassata (Hudson) Hazen
[
= Ch, endivcefolia Ag. Ok. Cornu Damce (Roth) Ag.], which possesses a tough,
;

gelatinous, sub-dichotomously branched thallus, rather flat and of a darker green

at the periphery than in the centre (fig. 27 A
and B). The branched thallus
bears much resemblance to the horns of a stag and in adult specimens is

frequently found floating freely at the marshy margins of ponds and lakes, or
even in bogs. Ch. tuberculosa (Roth) Ag. possesses a large cushion-shaped
thallus from 2 to 4 (or even 5) cms. in diameter, occurring usually in marshes
or in bogs. Ch. elegans (Roth) Ag. is a rarer species than the three previous
ones and occurs as very pale-green masses, clinging to submerged stems and
leaves of grasses, sedges, or mosses (fig. 27 C).

Genus Myxonema Fries, 1825. [Stigeoclonium Kiitz., 1843.]

The thallus is filamentous, branched, and usually devoid of the

great mass of gelatinous material

which is so conspicuous a feature
of Chcetophora. The branches are scattered, more or less isolated,
and often very elongated, but they are rarely developed in dense
fasciculate groups. The main branches bear other, shorter, lateral
branches which are either acuminate or terminate in long hyaline
hairs. There is usually a creeping portion of the thallus, attached
to some substratum, but adult plants frequently float freely in
1
of Ch. incrassata and Ch. elegans are often met with encrusted with
Forms
lime. A form
of the latter species was recently named by Tilden Ch. calcarea.
Vide Tilden in Botan. Gazette, 1897, pp. 97100, 102.
86 Chlorophycece

ponds and ditches. The macrozoogonidia and microzoogonidia

possess two cilia in some species, but four in others and they are
1
,

produced singly or in num-

bers from almost all the cells
of the thallus (fig. 28 B).
The gametes are biciliated
(fig. 280 and F) and the
zygospores are either smooth
or stellate. Iwanoff states
that the macrozoogonidia
germinate but that
directly,
the microzoogonidia pass
into a resting stage.
Fritsch 2 has recently
published some interesting
observations on early stages
of development of this genus.
He finds the development of
the basal portion to vary very
much in different species and
also to some extent within
the limits of each species.
He also concludes that cer-
tain plants described under
the generic name Herpo-
steiron are merely stages in
the life-history of epiphytic
Fig. 28. Myxonema tenue (Ag.) Rabenh., "
from near the Lizard, Cornwall. Stigeoclonia."
A, part of
thallus ( x 100) B, escape of zoogonidia; C,
;
Hazen 3 has given full
escape of gametes; D and E, zoogonidia; F, and conclusive evidence that
conjugation of gametes; G, development of a
cluster of zoogonidia ( x 500).
Myxonema was well estab-
lished before the publication
of Klitzing's genus Stigeoclonium, and sentimental reasons cannot
therefore stand in the way of the abandonment of the generic
"
name Stiyeoclonium."
There are several British species of this genus, of which the most frequent
is M. tenue (Ag.) Rabenh. (tig. 28). M. amcenum (Kiitz.) Hazen is also
another widely distributed species.
1
Iwanoff in Bull. Soc. Imp. Nat. Moscou, 1899.
-
Fritsch in Beihefte zum Botanischen Centralblatt, 1903, Bd xiii, Heft 4.
3
Hazen in Memoirs Torr. Bot. Club, 1902, xi, no. 2, pp. 193 4.
 ChcetopUoracece
Genus Draparnaldia Bory, 1808. The thallus is very gelati-
nous, and is differentiated into a principal filament and clusters of
lateral branches. The cells of the main filament are large, more
or less barrel-shaped, and are furnished with an equatorial, parietal

chloroplast with toothed edges. The main lateral branches are

alternate, opposite or verticillate, and are themselves very much

Fig. 29. Draparnaldia fllomerata (Vaucb.) Ag., from Tintagel, Cornwall. A, por-
tion of thallus ( x 100) ; B, single cell of main filament showing the chloroplast
(
x 220) ; C, part of branch showing escape of zoogonidia x 500)
( ; D, Ivypno-
spores formed from cells of branches ( x 500).

branched, the apical cells frequently terminating in long hyaline

hairs. From 1 to 4 zoogonidia arise in each cell of the lateral
branches (fig. 29 C) and they are furnished with four cilia. They
the cell-wall much smaller
frequently escape through a hole in
 88 Chlo rophycece
than their own diameter, and
they pass through many different
shapes in accommodating themselves to this small aperture. Usually
all the cells of a
single cluster of branches produce zoogonidia
simultaneously, the entire performance occu-
pying only a few minutes. Resting akinetes
(hypnospores) are frequently produced from
the cells of the branches.

There are two species, D. plumosa

(Vauch.) Ag.
and D. glomerata (Vauch.) Ag. (fig.
29), widely dis-
tributed in the British Islands.
They prefer clear
water and occur both in the still water of
bogs and
amongst stones in streams. When growing in
streams they are usually found in quiet pools,
stretching in long, pale-green, gelatinous strands
(up to 20 cms.) from stone to stone. The lateral
branches of D. glomerata are fewer and much
shorter than those of D. plumosa, the cells are
proportionately a little longer, and the hairs are
also usually longer. The diameter of the primary
filaments is 40 50 /x and that of the cells of the
branches 5 10 /*. Species of this genus are
amongst
the prettiest of all freshwater Alga?.

Genus Pseudochaete West & G. S. West,

1902. The thallus consists of two portions,
a creeping portion and an erect
portion. The
creeping part consists of cylindrical or barrel-
shaped cells, about 1^2^ times longer than
their diameter, each
containing a parietal
chloroplast with one pyrenoid. The erect
branches arise at right angles to the creeping
portion; they are narrower and attenuated
to fine points, each one
consisting of from
These cells are
five to eight distinct cells.
from 8 to 18 times
longer than their
diameter and each contains an elongated
Fig. 30. Pseudochcete
gracilis West & G. S. chloroplast, usually without a pyrenoid.
West, from near Coates, Sometimes the terminal cells of the branches,
Gloucestershire x 520).
(
or even the two terminal cells, contain no

protoplasmic contents and therefore no chloroplast.

P. gracilis West & G. S. West (fig. 30) occurs as an epiphyte on

aquatic plants. The diameter of the cells of the creeping filaments is 57
 Microthamniacece 8J
(

77 /x and of the erect branches only T5 1'8 p.. The plants bear considerable
resemblance to Herposteiron Nag. (Aphanochcete A. Br.) but differ in the
possession of completely septate branches instead of simple empty bristles.
The only other species of the genus is P. crassisetum West & G. S. West
which has been found in Ceylon, but it is quite possible that this is merely a
developmental stage of an epiphytic Myxonema.

Genus Thamniochaete Gay, 1893 The plants of this genus

1
.

are exceedingly minute, consisting only of three

to six cells. They are epiphytic and the basal
cell is
usually modified to form a hapteron.
The terminal cell possesses an elongated bristle
or a short spine-like projection. The chloro-
plast is
parietal and contains one pyrenoid.
Th. aculeata West & G. S. West is a rare plant
occurring as an epiphyte in the thallus of Gloeotrichia
natanx. It is only known from Connemara in Ireland
and from the Hebrides in Scotland. The diameter of
the cells is from 5'5 13 /z, and the terminal bristle is
short and very sharp, arising below the apex of a Fig. 31. Thamnio-
swollen terminal cell (fig. 31 A and B). Th. Hubert
Gay is epiphytic on a species of Oscillatoria and is from near Balallan
only known from the neighbourhood of Montpellier in Outer Hebrides B, ;

France from Baheh Lough,

rr,, , Galway. Ireland ( x
I his genus represents the simplest type of all the 520)
Chsetophoracese a type in which branching is practi-
cally absent. In Th. Hubert the terminal cells are attenuated and are
furnished with hollow bristles or hairs exactly as in many of the more
typical and complex Choetophoracese.

Family 6. MICROTHAMNIACE^l.
The thallus is filamentous, branched, and of small size. The
branches are never attenuated into hairs and the cells are some-
times moniliform or torulose. The chloroplast is a parietal plate
with one or many pyrenoids, or sometimes entirely without them.
The zoogonidia are only produced in special swollen cells of
the thallus which are differentiated as zoogonidangia. Reproduc-
tion frequently takes
place by akinetes.
It is a small family and bears considerable resemblance to the

ChsetophoracesB, being distinguished by the absence of multi-

cellular hairs and by the restricted origin of the It
zoogonidia.
1
Gay in Bull. Soc. hot. France, torn, xl, 1893, p. clxxvii cum fig. xylogr. 2.
90 CJdorophycece
also resembles the Trentepohliacese but differs in the aquatic
habit, smaller size, and in the nature of the chloroplasts.
Chodat includes the plants of this family in the Pleurococcacese.
but they have unquestionably reached a higher stage of develop-
ment than Pleurococcus or Trochiscia.
Genus Microthamnion Nag., 1849 ; Kirchn., 1878. The plants
of this genus are at first fixed but afterwards they often float

Fig. 32. A D, Microthamnion Kiitzingianum Niig. A C, young forms from

Richmond Park, Surrey (xoOO). D, portion of adult form from Horton-in-
Ribblesdale, W. Yorks. (x350). E, M. strictissimum Rabenh., from Blubber-
houses, \V. Yorks. ( x 500).

freely. The filaments are branched and the branches may be short
or long. The cells are cylindrical, 3 7 times longer than their
diameter, and the terminal cells of the branches are obtuse or
 Mia 'otham u iacece 91

acuminate. The branches all arise immediately below a trans-

verse cell-wall, and at first appear as lateral outgrowths from the

upper end of a cell. There is also a marked tendency for the

branching to be unilateral. The parietal chloroplast is long, entire,
and occupies about two-thirds of the inner wall of the cell it con- ;

tains no pyrenoids.

There are two species, M. Kiitzingianum Nag. and M. strictissimum Rabenh.,

1863 [
= J/. vexator Cooke, 1882]. The former species (fig. 32 A D) is much
more abundant than the latter, and is most abundant in the early spring. It
occurs in small pools and ditches, and likes peat. It is a small, much branched
plant, with short branches of one to six cells, the diameter of the branches
being 3 5 p.. M. strictissimum (fig. 32 E) is a larger plant with an erect
thallus up to 4 mm. in height. The branches are much longer and more
rigid, giving the plant a very different appearance from M. Kiitzingianum.
The diameter of the cells is 4 /^. Fig. 32 E is drawn from one of the original
specimens sent by W. B. Turner to M. C. Cooke when the latter described
the plant as "M. vexator."

Genus Gongrosira Klitz., 1843 [inclus. Pilinia Kiitz. (in part)].

The thallus is attached to a substratum by a mass of parenchyma-
tous cells formed by a confluence of creeping branches. From this
mass of cells, which may be one or many layers of cells in thick-
numerous, erect, branched filaments, varying in height
ness, arise
from 0'04 to 2 mm. This dense, cushion-like mass of erect fila-
ments is
frequently incrusted with carbonate of lime, and sometimes
forms quite a hard stratum. The cell-walls are often thick and
distinctly lamellose. The chloroplast is a parietal plate with one
or manypyrenoids, but it is usually difficult of observation. The
cellsgenerally present the appearance of being filled with a dense
chlorophyllaceous mass, which has been proved to contain starch
other than that present in the pyrenoids. The zoogonidia arise in
terminal zoogonidangia which are generally flask-shaped. The
akinetes are ordinary cells, generally of the recumbent portion of
the thallus, which become detached and ultimately form new
plants. The plants usually occur at the margins of ponds, lakes,
or rivers, forming a tough green stratum on submerged stones or
on the shells of aquatic Gastropods.

Species of this genus are rare in the British Islands, or perhaps they may
have been overlooked. G. viridis Kiitz. is a small species (thickness of prim. fil.
fig. 33 A C) usually encrusted with lime.
8 12 /i, of branches 4 8 ^ ;

G. stagnalis (West) Schmidle is a larger species (thickness of prim. fil.

92 Cliloropliycew

16 30 p.; fig. 33 D F) occurring attached to the shells of Limneea peregra.

Schmidle has recently given a short systematic account of the genus 1 .

Fig. 33. AGongrosira viridis Kiitz. (x500). A, from rocks, Lough Beg,
C,
Ireland B and
C, from rocks near Trernethick, Cornwall.
;
D F, G. stagnalis
(West) Schmidle, from near Button, Cambridgeshire ( x 200). zg, zoogoni-
dangium.

Genus Leptosira Borzi, 1883

2
This genus is scarcely to be
.

distinguished from Gongrosira. The plants are aquatic, forming

very minute bright green cushions. The branches are torulose, N
the terminal cells being elliptical, doliform, or sometimes irregular
in form. The cell-contents are pale yellow-green in colour and
their structure is exceedingly difficult to observe. The zoogoni-
dangia are intercalary and usually consist of the modified, older
cells of the plant. The zoogonidia may germinate directly or they
1
Schmidle in Berichte Deutsch. Botan. Gesellsch. 1901, Bd xix.
2
Borzi, 'Studi Algologici I,' Messina, 1883.
 Trentepohliacece 93

may conjugate in pairs and produce resting hypnospores. In the

conjugation of the zoogonidia the ends which do not bear the cilia
first fuse together.

L. Jfediciana Borzi is a rare plant found amongst Sphagnum and Utri-

cularia in bogs and boggy pools. It is only known from Yorkshire and
Sicily. Diameter of cells up to 20 p..

Family 7. TRENTEPOHLIACE^.
This family is only represented in the British Islands by a
few species of the genus Trentepohlia. The thallus is aerial,
filamentous and branched, generally occurring on rocks or on the
bark of trees. The filaments may be very short and more or less
creeping, or they may form erect tufts or closely matted cushions.
The sometimes cylindrical and sometimes moniliform or
cells are

torulose, and the branches usually show a slight attenuation. The

cell- walls are firm and
frequently exhibit external sculptures.
Brand states that the longitudinal walls are lamellose, but the
1

transverse walls are simple and that the cellulose caps which are
;

so frequently developed at the extremity of a branch, are the

remains of dead, terminal zoogonidangia. This is certainly
not true of some species, however. Each cell contains one
nucleus and usually a number of disc-like, parietal chloroplasts
without pyrenoids. The colour of these plants is usually some
shade of brown, brownish-red, or orange-red, the chlorophyll being
masked by the presence of a pigment known as hsematochromin,
which is frequently dissolved in a quantity of oil.
In the Trentepohliacea? the zoogonidia are only produced in
specially differentiated cells or zoogonidangia. This character, the
absence of terminal hairs and the nature of the chloroplasts, are
the principal distinctions between the TrentepohliaceaB and the
Chaetophoracese. The zoogonidangia are developed singly or in
clusters, either terminating a branch and so arresting its develop-
ment 2 or developed laterally on the branches, or more rarely in
,

'the axil of a branch. They are sessile or stalked, generally

ellipsoid or ovoid in shape, and they open by
means of a terminal
or subterminal pore. The zoogonidia, which are of two sizes, are
pear-shaped and furnished with two cilia. The smaller micro-
zoogonidia have been observed to conjugate in pairs, but all are
1
F. Brand in Beihefte z. Bot. Centralbl. xii, 1902.
2
The development is only arrested temporarily as the terminal cell or supporting
cell often grows through the empty zoogonidangium.
 I Chloropht/cece

capable of direct germination. Resting-spores or hypnospores are

sometimes produced.
Wildeman has shown the
1
ease with which these plants repair

injuries to the thallus.

Species of this genus are most abundant in damp tropical or
subtropical climates, occurring profusely as epiphytes on the leaves
and bark of trees. A
few of them are constituents of certain
tropical lichens.

Fig. 34. A C, Trentepohlia aurea Mart., from Cookridge, W. Yorks. ( x 500).

D F, T. calamicola (Zell.) Be Toni, from trees near Lough Gartan, Donegal,
Ireland ( x 500). zy, zoogonidangium.

]
Wildeman, in Mem. courormes et autres Mem. Acatl. roy. Belgiqne, 189!),
torn. Iviii.
 Uleacece 95

Genus Trentepohlia Martins, 1817. [Chroolepus Ag., 1824.]

The thallus is filamentous, simple or ramified. The branches are
alternate, of the same diameter as the principal filaments, and do
not terminate in a point or hair. The chloroplasts are numerous,
discoidal, and without pyrenoids, but are generally masked by
the presence of a red or orange-red oil which colours blue with
iodine.

The most abundant species in the British Islands is T. aurea Mart., which
occurs principally in hilly and mountainous districts, forming broad expanded
sheets of a bright red or orange-red colour. It is chiefly found attached to

rocks, particularly carboniferous limestone or silurian rocks, and generally

on
the windward side. The filaments are 1020 in thickness (fig. 34
//. C). A
T. odorata (Ag.) Wittr [=T. umbrina (Kiitz.) Bornet] and T. calamicola
(Zeller) De Toni (thickness of filaments 7'5 10 p; fig. 34 D F) are much
smaller British species of the genus.

Order III. ULYALES.

This order is mainly distinguished from the Chsetophorales by
the expanded, parenchymatous thallus, which is attached when
young to a substratum by rhizoids.' The cells are uninucleate
'

and they contain a single parietal chloroplast, often of considerable

bulk and containing one pyrenoid.
Asexual reproduction takes place by zoogonidia and also by
gemmation. Sexual reproduction is by isogamous planogametes
with two cilia.
There is only one family which has few freshwater repre-
sentatives.

Family 1. ULVACE^.
The Alga? belonging to this family are more often marine or
brackish in habit than freshwater. They consist of flat, ribbon -

plates, or more rarely they

exhibit a vesicular
shaped or expanded
or intestiniform structure. These flat or tubular structures consist
of one (Monostroma} or two (Ulva) layers of cells which may be
somewhat scattered and rounded in form, in which case they are
frequently arranged in groups of four,
or they may be closely
compact with polygonal outlines. The two genera Monostroma
and Enteromurpha have freshwater representatives, and in each
case the thallus consists of a single layer of cells, division only
are usually compact and
taking place in one plane. The cells
96 Chlorophi/cece

arranged with their long axes at right angles to the plane of the
thallus (fig. 35 D). Each cell contains a single nucleus and one
large parietal chloroplast, often with deeply incised or lobed margins
and containing a single pyrenoid. In Monostroma bullosa (Roth)
Wittr. the cells are generally arranged in T-shaped groups of four.
Asexual reproduction takes place in Ulva by zoogonidia (some-
times termed megazoospores) with four cilia. These come to rest
and germinate directly. Monostroma has been observed to re-
produce itself asexually by budding off small flat portions from
the surface of the thallus, each portion producing a new plant.
Geddes 1 has observed a process of gemmation in Enteromorpha.
Sexual reproduction is brought about by the conjugation of
isogamous gametes. Ordinary cells of the thallus become game-
tangia and give rise to eight (sometimes four or sixteen) piano-
gametes, which are pear-shaped bodies, smaller than the zoogonidia,
with a pigment spot and two long cilia. On conjugation the two
"
gametes coalesce slowly and a zygozoospore," or a rounded cell
with two pigment spots and four cilia, is first formed ;
this loses
its cilia and becomes a zygospore (fig. 35 J). The zygospore
usually germinates directly, forming a short filament of four
first

cells, which soon produce a flat expansion by dividing in two

directions in the same plane. According to Reinke the zygospore
sometimes becomes a resting-spore or hypnocyst, which on germi-
nation divides into four and then eight cells arranged peripherally
round a central cavity. By the increase of these peripheral cells
is produced, which in most instances
a vesicular thallus ultimately
becomes a flattened expansion attached by a few rhizoids at
its base.
Genus Monostroma Thur., 1854. The thallus in the adult
is always a thin membranaceous plate. In its younger
plant
stages it is frequently vesiculose, opening out as
grows into a it

foliaceous plate, finally becoming free-floating. It consists of a

single layer of rounded or more or less angular cells which are

often disposed in groups of four. The zoogonidia possess either
two or four cilia, and the gametes are biciliated and rather smaller
in size.

Few species of the genus inhabit fresh water, the only British representa-
tives being M. bullosa (Koth) Wittr. and M. membranacea West & G. S. West

1
Geddes in Trans. Roy. Soc. Edinburgh, 1881, p. 555.
 Ulrae-ew 97

(fig. 35,
A
K). In the former species the cells are rounded, 6 12 // in
diameter, and arranged in fours, the two pairs often being disposed in a more
or less T-shaped manner. In the latter species the cells are much more
compact, angular, and 8 20 p.
in diameter.

Fig. 35. A K, Monostroma membranacea West & G. S. West, from Mitchain

Common, Surrey. A, nat. size B and C, portions of thallus showing cells
; ;

I), section of thallus; E, cells with escaping gametes (x566). F J, con-

jugation of gametes (F H, x 566; I and J, x 790). K, young plant developed
from hypnospore (x566). L, Enteromorpha intestinalis (L.) Link, from
Frizinghall, W. Yorkshire (nat. size).

Genus Enteromorpha Link, 1820. The thallus is elongated,

tubular and intestinifurm, sometimes reaching a considerable
length. It is green, yellowish-green, or pale olive-green in colour,
and consists of a single layer of rounded or polygonal cells, each
with a parietal chloroplast.

The only freshwater species of the genus is E. intestinalis (L.) Link

(fig.35 L), an Alga which also occurs in brackish water and in the sea. It is
widely distributed in the large drains and dykes in the east of England, and
also occurs frequently in cana.ls and ponds in other parts of the country.

W. A.
98 Chlorophycefc

Order IV. SCHIZOGONIALES.

The thallus filamentous, sometimes (especially in young
is

stages) parenchymatous, and often expanded into broad sheets by

the fusion of the filaments in one plane. The cells are unirucleate
with a single central, stellate chloroplast, containing one pyrenoid.
The plants are often attached by 'rhizoids' to a substratum,
and are subaerial in habit.
The order at once distinguished from the Chastophorales by
is

its chloroplasts,and by the division of the cells in two, and often in

three directions, especially in young plants. From the Ulvales it
is distinguished by its chloroplasts, by the more or less regular

longitudinal arrangement of the thallus-cells, and by the absence

of the vesicular stage in the growth of the young plants. The
plants of this order have most probably had a very different origin
from the Ulvales, the resemblance being only a parallelism of
modification.

Family 1. PRASIOLACE^El.
This family has been established to include those Alga3 which
are embraced in the Schizogoniales.
The thallus is commonly terrestrial, simple and filamentous, or /
forming flat, creeping expansions. It consists of a single layer of
cells produced largely by a fusion of the contiguous walls of cell-
filaments. Each cell possesses a central, stellate chloroplast with
one pyrenoid.
Asexual reproduction takes place by gemmation and by the
formation of resting akinetes. Lagerheim has observed the pro-
'
duction of tetraspores.'
Chodat regards the family
as having analogies to the Bangiacese

among the Rhodophycere, both on account of the production of

tetraspores and the mode of growth.
Genus Prasiola Ag., 1821. [Indus. ScJiizogoiiiam Klitz. 1843,
and Hortniditim Kiitz. 1843.] The genus occurs on moist earth,
rocks, stones, old walls, trunks of trees, etc. The thallus is
filamentous, consisting of one, two, or many series of cells, or
foliaceous and expanded with the cells arranged more or less in

groups of four. The cells of the ordinary filaments are broader

than long and those of the flat expansions are quadrate or polygonal
 Prasiolacece 99

in form. The and hyaline. The single

cell-wall is strong, rigid

chloroplast central, star-shaped, and contains one pyren<>id.

is

Sometimes the thallus is fixed by rhizoids and sometimes not. In

the broader, flat expansions the cell-walls are thick and confluent,

Fig. 3G. A C, Prasiola pariettna (Yauch.) Wille, from Bradford, W. Yorkshire

(
x 500). D G, Prasiola crispa (Lightf. Menegh. D, examples from Bradford,
) ;

W. Yorks. (nat. size) E, simple filament from Helvellyn, Westmoreland

; ;

F, portion of irregular filament from Wimbledon Common, Surrey ( x 500) ;

G, basal portion of broader thallus, from Bradford, W. Yorks. ( x 400).

and the cells have the appearance of being separated by consider-

able spaces. Reproduction is by a process of gemmation, by
akinetes liberated at the margins of the thallus, and by tetraspores 1 .

Gay and Chodat separate the genera Schizogonium and Prasiola,

but I am inclined to agree with Wille 2 in uniting them under
Prasiola Ag.
3
Wille and Borgesen have each described some interesting
marine forms of this genus in which the plants are more amply
supplied with rhizoids. The expanded thallus of these forms does
not reach such a large size as the thallus of the land forms.

Two species are abundant, P. crispa (Lightf.) Menegh. [which includes

Hormidium murale Kiitz.; Schizogonium crispum (L.) Gay; and Ulothrix
radicans Kiitz.], the cells of which are 7 14 /x in diameter (fig. 36 D G),
and P. parietina (Vauch.) Wille [which includes Schizogonium murale Kiitz.
and Hormidium parietinum Kiitz.] with cells 9 18 /z
in diameter (fig. 36
T
Lagerheim, Ueber die Fortpflanzung von Prasiola,' Ber. Deutseh. Botan.
'
,

Gesellsch. 1892, Bd x, Heft 7.

2
Wille, 'Studien iiber Chlorophyceeu I- VII,' Videusk. Skrifter, I math.-
naturv. Klasse, 1900, no. 6, p. 13.
3
Borgesen, Marine Alg. of the Faeroes,' Bot. of Faeroes, Part II, 1902.
'

72
100 Chlo 'ophycece
i

A C). These two species are widely distributed all over the British Islands,
and they have a decided preference for the neighbourhood of towns, being
found frequently under walls and as a green carpet between the paving-stones
of quiet streets. They require little moisture and can withstand considerable
desiccation. Associated with them are generally numerous Rotifera mdgaris
and testaceous Rhizopods such as Trinema acinus. Prasiola furfiiracea
Menegh. is probably a form of P. crispa.

Order V. MICROSPORALES.
This order was first established by Bohlin to include those
curious plants which belong to the genus Microspora. It seems
at first sight to be giving undue prominence to a small
group of
aberrant Algas, but at the same time it removes a difficulty, as
these plants cannot well be placed in any of the other orders of
green Algas.
The thallus is filamentous and unbranched, and the cell-walls
frequently become broken up into H -shaped pieces. The cells are
uninucleate, with a large reticulated chloroplast occupying almost
the entire inner surface of the cell- wall, and destitute of pyrenoids.
The affinities of the order are very doubtful.

Family 1. MICROSPORACE^.
This small family includes only one genus. The thallus is
filamentous and simple, and the cells are cylindrical. The cell-
walls are composed of cellulose, are either homogeneous or more or
less distinctly lamellose, and of a similar structure to those of
Tribonema (Conferva), the cells often becoming disarticulated into
H -shaped pieces. A single nucleus of considerable size is
present
in the centre of each cell. The
chloroplast is disposed on the walls
of the cell and may be band-like or sheet-like, covering more or less
the entire cell-wall. It is usually areolated or reticulated, and
really consists of a fusion of numerous cushion-like, chlorophyl-
laceous masses to form a stout areolated structure. There are no
pyrenoids, but scattered granules of starch are often present.
Asexual reproduction takes place by the formation of aplano-
spores which become hypnospores (fig. 37 C and F) also by the ;

production of biciliated or quadriciliated zoogonidia, one or two

of which are found in a cell. Sometimes several small micro-
zoogonidia are produced in a cell. The zoogonidia germinate
directly.
 Micro ^ o ) i 'acece 101

Genus Microspora Thuret, 1850; em. Lagerh., 1888. The

filaments are simple, con-
sisting of cylindrical or
slightly swollen cells. The
cell-walls are firm, fre-

quently distinctly lamel-

lose, and sometimes dis-

sociating into pieces

which appear H -shaped
in optical section, each
piece consisting of a trans-
verse wall and portions
of the lateral walls of two

adjacent cells. There is

one cell-nucleus. The

is disposed on
chloroplast
the cell-walls as a more
or less reticulated mass.
Globular hypnospores are
produced, with thick walls, 37. A, Microspordamc&na (Kiitz.) Lagerh.,
.

from near Senens, Cornwall. B and C, M. nb-

usually one in each cell. brerinta (Babenh.) Lagerh.; B from Tremethick
Moor and C from St Just, Cornwall. D, M.
There are several widely
pachyderma (Wille) Lagerh., from near Land's
distributed species of this End, Cornwall. E and F, forms of M. anni'ini
genus in the British Islands. (Kiitz.) Lagerh.; E, from Shipley, W. Yorks.,
to show the chloroplast; F, from New Forest,
M. floccosa (Vauch.) Thur.
Hants, n, aplanospores. (All x520.) E is
and M. amosna(Kutz.} Lagerh. 31. ama'tia var. crasnior Hansg.
(fig. 37 A, E and F) are the
most abundant and often occur in small ponds and horse-troughs. M. abbre-
viata (Rabenh.) Lagerh. (tig. 37 B and C) and J/. pachyderma (Wille) Lagerh.

(fag. 37 D) are not so frequent.

Order VI. CLADOPHORALES.

In this order are included three families of green Alg which
bear close relationship to the Siph'onese and yet can scarcely be
relegated to that order. The thallus is simple or branched and
incompletely septate, each segment containing many nuclei and
numerous parietal chloroplasts, the latter containing single
pyrenoids.
Asexual reproduction takes place by biciliated or quadriciliated
zoogonidia, by 'cysts,' or by special resting-spores (Pithophora) ',
102 Chlorophyeece

and sexual reproduction is either by isogamous planogametes or

by well-differentiated heterogamous gametes.
The order includes the following three families :

Family 1.Cladophoracece. Thallus large, branched, incompletely

septate, usually attached. Segments large with numerous small
chloroplasts, each with a pyrenoid.
Family 2. Pithophoracece. Thallus similar to that of the Clado-
phoracese, but distinguished by the formation of barrel-shaped and
fusiform asexual resting-spores.
Spkoeropleacece. Filaments simple, composed of elongated
Family 3.
ccenocytes. Sexual reproduction heterogamous plants monoecious
; ;

oogonia with many non-motile oospheres fertilization within the

;

oogonium.

Bohlin has recently proposed to transfer the families Clado-

phoracese (including the Pithophoraceae) and Sphasropleacea? to
the order Siphoneae, placing them next the Yaloniacese. In this
'he is followed by Blackman and Tansley, but it yet remains to

be shown how far this change is justified. The thallus of the

Cladophorales is much more septate than that of the Siphoneas

and the branching is of a different nature.

Family 1. CLADOPHORACE^I.
The thallus is large, filamentous and incompletely septate, each
segment being a ccenocyte. The filaments usually have a basal
organ of attachment, and in the genus Cladophora are much
branched. The growth of the thallus is
apical in Cladophora but
intercalary in the other genera.
There are several nuclei in each segment of the thallus and
either one reticulate, parietal chloroplast or a large number of

separate chloroplasts, each with a pyrenoid.

Asexual reproduction takes place in Cladophora and Chcvto-
morpha by zoogonidia which are produced in large numbers in the
mother-cell and escape either by a terminal or lateral pore. 'Cysts'
are also produced in Rhizoclonium, each one being a kind of large
thick-walled akinete formed from a single segment.
An isogamous sexual reproduction occurs in Cladophora, the
conjugation of the planogametes resulting in a zygospore which
germinates directly without rest.
Very little, if any, mucus is secreted by these Algae and they
always feel rough and crisp. The absence of a mucous outer coat
 Cladophoracetr 103

causes them to be frequently loaded with epiphytes. Very often

the older filaments are thickly covered with Diatoms, particularly
of the genera Gomphonema and Gocconeis.
Genus Chaetomorpha Ktitz., 1845. The filaments are simple,
ofmore or less uniform thickness
and fixed at the base, the lower
segments being shorter than the
upper ones. The segments are
often slightly swollen and they
are of considerable diameter. The
cell-wall is thick, very firm and
obviously lamellose.
Most of the species of this genus
are truly marine or brackish in habit,
but Ch. sutoria (Berk.) Eabenh. (fig. 3S)
issometimes found in running water, in
we] Is and horse-troughs, or in streams.
Diameter of filaments 100 120 p.
Fig. 38. Cluetomorphasutoria (Berk.)
Genus Rhizoclonium Kiitz., Eabenh., from Heaton, W. Yorks.
(
x 100).
1843. The filaments are of vari-
able size, crisp, generally branched, and attached at the base by a
branched hapteron. The branches are short, slightly attenuated,
sometimes merely unicellular outgrowths, but more frequently
consisting of several cells. The filaments are often bent at the
point of origin of a branch. The cell-walls are firm, lamellose,
and sometimes attain a considerable thickness. The number of
nuclei present in each segment is variable and the chloroplast is
in the form of a network containing several pyrenoids. The most
recent account of the structure of this genus is by Wille
1
who ,

2 3
confirms much that has been described by Gay Stockmeyer .

has given a thoroughly good systematic account of the genus.

In the autumn the segments of the thallus are often packed
with starch.
or they may even
Species of this genus are marine, brackish, freshwater,
occur on damp soil. The only freshwater British species is R. hieroglyphicum
Kiitz.; em. Stockm. [= Conferva fontinalis Berk.; Microspora fonti'milis
De
1
Wille, 'Studien iiber Chlorophyceen VII,' Vid.-Selsk. Skrifter. M.-N. Kl.
Christiania, 1900.
2
Gay, 'Eecherches sur le devel. et la classif. de quelques Algues Vertes,' Paris,
1891.
Algengattung Rhizoclonium,' Verhandl. der k.
3
Stockmeyer,
'
Uber die k.

Zool.-Bot. Gesellsch. in Wien, Jahr. 1890.

104 ChlorophycecB

Toni] which possesses filaments 10 37 p in diameter and segments

2 5 times

longer than the diameter (fig. 39 A). The common form of

this species pos-
sesses no branches whatever
and occurs abundantly in ponds,
ditches, drains, streams and
cataracts all over the country,
B
thriving well in water in which
considerable putrefaction is

taking place. The following

varieties of it are widely distri-
buted : -var. Kochianum (Kiitz. )

Stockm. [=R.KochianumK\itz. ;

R. flavicans Rabenh.], var. tor-

tuosum (Kutz.) Stockm. (fig.
39 B E), and var. riparium
(Harvey) Stockm.
Genus Cladophora
Kutz., 1843. This is the
best known genus of the
family and is widely dis-
tributed in salt and fresh
Fig. 39. A, Rhizoclonium hieroglyphicum waters. The thallus is
Kiitz., single cell showing chloroplast and the of
from branched, type
pyrenoids, Sheep's Green, Cambridge
(x500). B E, E. hieroglyphicum Kiitz. var. branching varying with
tortuosum (Kiitz.) Stockm., from Heaton, different species, and the
W. Yorks.; B and C, x 100; D and E, x 500.
segments are 6 12 (or
even up to 20) times longer than their diameter. There are usually
many nuclei in a segment, but they may be reduced to two or even
but
one. The parietal and most commonly reticulate,
chloroplast is

all intermediate stages are met with between an elongated reticu-

late cylinder and isolated plates
1
There is one pyrenoid in each
.

isolated plate or in each corresponding piece of the reticulum.

The cell -wall consists of an inner and outer layer, and, according
to Brand, of an 'outermost' layer which can be separated by acetic
acid. The zoogonidia are
very numerous and escape from the
mother-cell through an opening formed by a complete absorption
of the cell-wall. Nordhausen 2 regards the basal branching of the
to
segments in a filament of Cladophora as a peculiar process,
which there is nothing strictly comparable in other Algse. Brand
states that the species with strong, primary, basal organs of attach-

1
Brand in Beitr. z. Bot. Centralbl. x, 1901.
2
Nordhausen in Pringsheim's Jahrb. f. wiss. Bot. xxxv, 1900.
 Cladophoracece 105

ment usually form zoogonidia, and that the free-floating species

are usually propagated by resting-spores.

Fig. 40. Cladophora glomerata (L.) Kiitz., from Shipley, W. Yorks.

A, nat. size; B, x 85.

Most of the species of the genus are marine, but some five or six British
freshwater species are known. Cl. glomerata (L.) Kiitz. (fig. 40) is an abundant
species, occurring as dark green masses attached to rocks and stones in
streams and waterfalls. The branching is dense and the smaller branches
are in tufts. Cl. crispata (Roth) Klitz. is another common species, usually

occurring attached to stones. Cl. flavescens Ag. is a more slender species of a

pale yellowish-green colour and frequently occurs floating freely in ponds and
106 Chlorophycece
ditches. Brand 1 states that
allthe European species of
Cladophora described by Ra-
benhorst, except the JSgagro-
pilse,must be regarded as
varieties, forms, or conditions
of (.7. fracta or CL glume-
rata.

Genus Chsetonella
Schmidle, 1901-. This
genus was instituted
for the reception of a
small, almost microscopic
plant, which occurs either
free-floating or attached
to the mucous coat of
other larger Algae. It is

incompletely septate, and

each segment possesses
from two to five nuclei.
The segments are cylin-
drical or more or less ir-

regular, and the thallus is

branched. The branches

are attenuated and the
terminal cells are of

greater length than the

others. There is one
parietal chloroplast with-
out pyrenoids.
There is only one species,
Ch. Goetzei Schmidle, which is
Fig. 41. Pithophora (Edogonia Wittr. var. known from tropical Africa
pohjitpora Rendle and West f., from the Eeddish
Canal, near Manchester, A, x 65 B D, frag-;
and from West Yorkshire.
ments of thallus with asexual resting spores (as), The filaments are 6 8 p in
xlOO. thickness.

Family 2. PITHOPHORACE.EJ.
The
plants of this family very much resemble species of the
genus Cladophora. The thallus is of exactly the same type and the
1
Brand in Bot. Centralbl. Ixxix, 1899.
2
Schmidle in Engler's Botan. Jahrbiich. Bd 30, Heft 2, 1901, p. 253, t. v,
f. 1, 2.
 Pithophoracece 107

growth is apical. The thallus is almost always branched, the

branches arising a little below the top of their supporting cells,
and it is attached below by well-developed haptera put out from a
basal cell. It is doubtful if there is sufficient evidence to warrant
the separation of the Pithophoracese as a distinct family from the
Cladophoracese.
Asexual resting-spores are produced here and there in the
thallus, either intercalary and cask-shaped, or terminal and ovoidal
or fusiform. These spores, which were termed by Wittrock 'agamo-

spores,' are richly filled with chlorophyll and the spore-wall increases
considerably in thickness. Formation of spores may take place in
allthe cells of the cauloid part of the thallus. After a short period
of rest the spores develop in opposite directions from the two
'

apices. There is another method of asexual reproduction by pro-

lific cells.'

Genus Pithophora This striking genus of Algse

Wittr., 1877.
is almost exclusively tropical in its distribution. It is the only

representative of the family, and few Algas are more characteristic

than a plant of Pithophora with ripe spores.

P. (Edogonia (Mont.) Wittr., var. polyspora Rendle and West f. 1 (fig. 41)

may be regarded as a British Alga, having been found in the Reddish Canal,
near Manchester. P. Keivensis Wittr. occurred in a tank in the water-lily
house, Kew Gardens, but was most probably introduced from tropical
S. America.

Family 3. SPH^ROPLEACE^.
This family includes only one Alga, which occurs in extensive
masses on flooded plains and by the margins of lakes. The
filaments are cylindrical and imbranched, and consist of single
series of coenocytes which reach an extraordinary length as com-

pared with their breadth. The transverse cell-walls often become

of great thickness, and each segment of the filament contains a
considerable number of small nuclei. The chloroplasts, which arc
in the form of parietal rings, are very numerous and some of them
contain pyrenoids.
The sexual organs are oogonia and antheridia, which may be
formed without change of shape from any segment of the filament.
Sometimes the oogonia and antheridia alternate in a filament, but
1
A. B. Eendle and W. West, jtmr. ,
'A New British Freshwater Alga,' Journ.
Bot. July, 1899, pp. 289291, t. 399.
108 ChloropTiycece

more often they do not. A number of oospheres are developed in

each oogonium, and a very large number of antherozoids arise by
the breaking up of the red contents of an antheridium. The
antherozoids are small, elongated bodies, provided with two long
cilia, and they find their way into the oogonia through apertures
in the transverse walls. After fertilization, which takes place
within the oogonium, the oospores develop a thick verrucose
cell-wall and the cell-contents become bright red in colour. The
oospores generally hibernate within the oogonium, and this is the
most striking condition of the Alga, Avhich now consists of long
filaments, most of the segments of which are filled with numerous,
bright red, verrucose oospores.
On the germination of the oospore from two to eight zoospores
are set free, representing the sporophyte generation, and each of
these forms a new plant. The young plants are simple, fusiform
cells, with each extremity attenuated to a very fine point.
Bohlin has suggested the transference of this family to the
order Siphonese in close proximity to the family Valoniacese.
Genus Sphseroplea Ag., 1824. The thallus consists of un-
branched filaments of cylindrical coenocytes, which may only be as
long as their diameter or up to ninety times longer. The filaments
are 36 72 a in diameter.
The only known species Spk. annulina (Roth) Ag. is not a British
Alga. It occurs extensively on inundated portions of the plains of Europe,
Asia and America, and is sometimes found in pits or quarries. It is question-
able there are any suitable localities for this plant in the British Islands.
if

It sometimes occurs in Kew Gardens, having been introduced from abroad

with various aquatic plants.

Order VII. SIPHONED.

The order Siphoneas embraces a large number of filamentous
Alga3 of many diverse forms. They are coenocytic in character,
the individuals being without septa, so that in reality each plant
consists of a single large coenocyte. They are mostly marine Algre
and some of them attain a considerable size. The thallus fre-
quently becomes very complicated in character, but even then it
consists of the interlaced branches of a single multinucleated cell.
Nowhere else in the vegetable kingdom do such gigantic cells

occur, and it is this extraordinary complication of a single cell

which distinguishes the Siphonea? from all other Algae. Many of

 Vaucheriacece 109

them have even reached a

high stage of specialization, having
developed organs analogous to the stem, leaf and root of higher
plants. In the genus Gaulerpa the cell is strengthened by trabe-
culsewhich traverse the lumen of the cell from wall to wall.
Asexual reproduction takes place by proliferous shoots, by
non-ciliated spores and by zoogonidia. In most of the Siphoneae
numerous zoogonidia arise in a zoogonidangium, but in Vaucheria
only one large one is produced.
The plants are generally attached by strongly developed haptera.
The only family of the order which inhabits fresh waters is the
Vaucheriacese, and it is also the only family in which well-
differentiated sexual organs occur. The tropical family Phyllo-
siphonacea? includes a number of AlgaB which live as parasites on
the leaves of Phanerogams.

Family 1. VAUCHERIACE.EJ.
The an elongated filament consisting of a single large
thallus is

coenocyte, and sometimes branched. This imseptate filament

is

increases in length by apical growth and is

usually attached to a
substratum by much branched haptera. In most members of the
family the cell-wall is thin and relatively weak, easily collapsing-
even with careful manipulation. The protoplasm forms a thick
lining layer on the interior of the wall of the filament and contains
a large number of minute nuclei. The chloroplasts are very small
and exceedingly numerous they are oval, elliptical or subcircular
;

in outline and are without pyrenoids. A considerable amount of

often present in the filaments, the oil-drops
oil is
being always in
connection with the chloroplasts. Fleissig 1 states that this
oily
material is a reserve substance, physiologically
analogous to starch.
On the injury of the thallus septa usually appear cutting off
the injured parts, the uninjured portions developing into new
plants, (vide fig. 42 A and B.) These are the only instances of
the occurrence of septa in the thallus except in connection with
the reproductive organs.
Asexual reproduction takes place by the formation of zoogonidia.
The extremity of a filament assumes a
club-shaped form and
becomes densely filled with protoplasm, after which a transverse
1
Fleissig, 'Ueber die phys. Bedeutung d. oelartigen Einschlusse in d. Vaucheria,'
Basel, 1900.
110 Chlorophyeece

septum appears and cuts off the swollen end as a zoogonidangimn.

The contents of this gonidangium, which are of a rich green colour,
gradually become rounded off, forming an oval zoogonidium of
large size. The whole surface of the zoogonidium is usually
clothed with numerous short cilia, arranged in pairs, and in the
surface protoplasm under each pair there is a small nucleus. The
entire structure contains one central cavity filled with cell-sap

Fig. 42. A and B, portions of thallus of Vauclieria showing formation of septa on

injury; A, Vaticheria geminata (Vauch.) D. C.; B, V. sericea Lyngb., from
Harerield, Middlesex. C, apex of filament of V. sessilis (Vauch.) D. C., show-
ing the zoogonidangimn from which will escape a single zoogonidium. D, ger-
mination of the zoogonidium of V. sericea, from E. Yorkshire. E, V. sesxili*
from W. Yorkshire, showing developing oogonium (on right) and antheridium
(on left). F H, V. geminata (Vauch.) D. C., from Barnes Common, Surrey,
showing development of sexual organs. (All x 75.) a, antheridium, oo, oogo-
nium; zg, zoogonidangium.

which is traversed by strands of protoplasm, and Schmitz has

pointed out that it can be looked upon as an aggregate of zoogonidia
with a great resemblance to certain of the more complex members
of the Yolvocaceae. This compound zoogonidium escapes by an apical
opening of much smaller diameter than itself and through which it
pushes its way. Sometimes, owing to the rotatory movement of the
cilia, the part first exuded becomes separated from the portion still
left in the gonidangium and two zoogonidia are formed instead

of one.
The zoogonidia generally escape in the morning, that is to say,
after the plants have been in the darkness for some time. They
 Vauclieriacece 111

are sluggish in their movements and continue active" for about

fifteen minutes. On coming to rest the cilia are at once withdrawn
and a cell-wall is
developed. Klebs states that zoogonidia can
1

always be produced when filaments which have been kept moist

for some days are soaked with water, or when they are removed

from a dilute nutritive solution into pure water. The zoogonidia

germinate almost immediately by the protrusion of one or more
tube-like filaments, one at least of which attaches itself to the
'
substratum by a colourless branched hapteron (or rhizoid ').
Under unfavourable circumstances, particularly if the plants
are liable to become dried up, asexual spores of another kind are
sometimes developed. The end of a filament swells up into a
more or less globular form and then a transverse wall appears and
cuts this portion off. Such a spore may rest a considerable time
before germination.
Sometimes, owing to drought, certain filaments break up into a
number of distinct ccenocytes, each of which develops a thick cell-
wall. These are of the nature of rudimentary gemma? or cysts 2 .

Sexual reproduction takes place by oogonia and antheridia, and

amongst Siphoneae this family of Algse stands alone in the
^the

possession of sharply differentiated sexual organs. These are

developed at scattered intervals along the cylindrical filament.
Except in the dioecious plants the antheridia and oogonia usually
arise sideby side on the same filament, or they are differentiated
portions of a short lateral branch.
The oogonia usually arise as lateral outgrowths of the filament,
or at the end of a very short branch, and they soon assume a more
or less rounded or ovate form, being ultimately cut off by a septum
at the base. The apex of the oogonium generally develops a
rostrum or beak, which is usually turned to one side, either towards
the antheridium or away from it. The protoplasm of the oogonium

1
Klebs, 'Die Bedingungen d. Fortpflanzung bei einigen Algen u. Pilzeu,' Jena,
1896.
2
Bennett and Murray in their 'Handbook of Cryptogamic Botany,' London,
1889, p. 284, in referring to this special type of asexual reproduction in Vaucherin,
state that " in this condition it was formerly described as a distinct organism under
the name of Gongrosiru." Dr Scott, also, in his Introd. to Structural Botany,
'

Part u,' London, 1897, remarks that "this is called the Gongrosira state, because
specimens of Vauchcria in this condition used to be placed in a different genus
under that name." These are most unfortunate statements, as Gongrosira is a
well-established genus of the Chaetophorales, which reproduces itself asexually by
minute biciliated zoogonidia, and none of the plants of which approach in size even
the smallest known species of Vaucheria.
112 CMorophycece
contains a considerable amount of oil, numerous chloroplasts, and
after the appearance of its basal wall, only one nucleus. That
portion of the protoplasm towards the apex or beak becomes clear
and free from chloroplasts, and is termed the receptive spot. The
cell-contents now become rounded off, forming the ovum (or

a-.

00

Fig. 43. Aand B, Vaucheria sessilis (Vauch.) D. C. A, from Esher West-end

;

Common, Surrey; B, from Mitcham Common, Surrey. C and D, V. hamata

(Vauch.) Lyngb., from Calverley, W. Yorks. E, oogouiurn and oospore of
V. sericea Lyngb., from Beverley, E. Yorkshire. (A D, x200; E, x320.)
a, antheridium; an, antherozoid; oo, oogonium.

oosphere) and soon afterwards the wall of the oogonium opens at

the extremity of the beak, a small quantity of mucilaginous pro-
toplasm being exuded.
The antheridia develop simultaneously with the oogonia and
generally in close proximity to them. (Fig. 42 E.) Each anthe-
ridium arises as a short cylindrical branch which usually becomes
much curved on approaching maturity. (Fig. 43 A and B.) The
terminal portion of this curved branch is cut off by a septum and
becomes the actual antheridium. In some species, such as in the
submarine Vaucheria synandra, a number of antheridia occur on a
"
structure known as an androphore." The protoplasm of each
antheridium contains numerous chloroplasts and nuclei. The
nuclei collect in the central portion of the antheridium and it is
 Vaucheriacece 113

this part which breaks up into the antherozoids (or spermatozoids).

The latter are extremely minute, each one consisting of a small
amount of protoplasm surrounding a nucleus and possessing two
cilia. The cilia are attached far apart and point in opposite
directions. (Fig. 43 A, an.) The antherozoids swarm for a short
time within the antheridium, which soon opens at the apex and
sets them
free. A certain amount of unused protoplasm is
expelled with the antherozoids and another portion is left behind
in the antheridium.
The antherozoids swarm near the opening of any oogonium

they may happen to reach and frequently several of them enter

the oogonium. One of them fuses with the oosphere at the region
of the receptive spot and its nucleus travels through the protoplasm
of the oosphere until it reaches the single nucleus of the latter.
The male and female nuclei then unite and fertilization is effected.
The fertilized ovum or oospore now invests itself with a cell-
wall of considerable thickness and undergoes a prolonged rest.
The oospores can withstand a certain amount of desiccation, and on
germination they give origin to a new plant without any alternation
of generations.
Genus Vaucheria DC., 1803. This is a widely distributed
genus of Alg?e the filaments of which are interwoven to form
compact, mat-like masses, either on damp earth, or in fresh or salt
water. They most commonly occur in situations where they are
subject to the splashing or trickling of water, although some
habitually occur on damp ground or are entirely submerged.
They
are most abundant in the earlier months of the year. The filaments
are coarse and thick, consisting of large ccenocytes, which some-
times reach 30 cms. in length, a.nd their numerous chloroplasts
give them a dark green colour. They are very sparsely branched,
and some species rarely exhibit any branching except in connection
with the formation of the sexual organs. Dioecious species of the
genus exist, but in most others the antheridia and oogonia are
developed in close approximation on the same filament. The
filaments are sometimes subject to the attacks of the Rotifer
Notommata Wemeckii, which produces irregular gall-like swellings.

V. sessilis perhaps the commonest species of the genus,

(Vauch.) DC. is

being widely distributed in damp and wet situations in the neighbourhood of

streams, cataracts and boggy springs thickness of filaments 65 80 p (tigs. 42
;

C and E 43 A and B). V. geminata (Vauch.) DC. also occurs in similar

;

w. A. 8
114 Chlorophycece
localities ;
thickness of filaments 78 90 /* (fig. 42 A, F H). V. terrestris

Lyngb., and 'Yauch.) Lyngb. (fig. 43 C and D), occur in profusion

V. 1m, ,,<//</
on damp ground, often forming thick mats on gravel paths and on the surface
T
soil of damp flower-pots. T sericea Lyngb., which is the smallest British
.

species (thickness of fil. 48 55 fig. 42 B and D, 43 E), and V. aversa Hass.

fj. ;

usually occur entirely submerged in the waters of ditches and ponds. V. dicho-
toma (Lyngb.) Ag. is the largest British species (thickness of fil. 180 220 p.)
and is dioecious. Some forms of it are truly marine, but others occur in
brackish water.

Order VIII. CONJUGATE.

The order Conjugate is one of the best defined and most
natural the groups of the Chlorophycese.
of The thallus is
unicellular in the Desmidiacese and the individuals exhibit great

specialization of form. In the Zygnemacese, which is the only

other family of the order, the plants are multicellular, consisting
of unbranched filaments of cylindrical cells. These filaments are,
however, fragile and often become dissociated into their individual
cells. All the plants of this order, whether unicellular or multi-
cellular, are remarkable for the great development of the gelatinous
pectose constituents of the cell-wall. There appears to be a
continual exudation of this gelatinous material, until, in some
instances, it is of much greater bulk than the individual plant,
and frequently happens that the unicellular forms occur
it

embedded in a mass of transparent jelly formed by the coalescence

of their outer gelatinous coverings.
One most conspicuous features of the order is the
of the

presence of chloroplasts of large size and definite form. They vary

in number from one to about eight or twelve in each cell and they
exhibit great variety in form and symmetry. Each' chloroplast
contains one or more conspicuous pyrenoids. Boubier 1 has
observed in species of Spirogi/ra and in Mougeotia scalaris Hass.
what he terms compound pyrenoids.' These consist of an
'

agglomeration of pyrenoid structures enclosed in a membrane and

containing in the centre a pyrenocrystal.
Multiplication of the filamentous forms sometimes takes place
by the fragmentation of the filaments, each cell undergoing rapid
division and forming a new plant.
Asexual reproduction may be brought about in the Zygnemacese

1
Boubier in Bull. Herb. Boissier, vii, 1899.
 ConjugatCB 115

by the formation of with thick walls, which have

resting-cells
received the namecysts.' These
of
'
are capable of surviving the
winter. In both the Zygnemaceee and Desmidiaceee aplanospores
are sometimes formed. Zoogonidia are entirely absent from this
order of green Algae.
Sexual reproduction takes place by the conjugation of isogamous
gametes and the formation of a zygospore. As the gametes are
devoid of cilia and therefore non-motile, they are known as aplano-
gametes. The ordinary vegetative cells become the gametangia,
usually without change of form, and only one gamete arises from a
gametangium, the entire cell-contents of which are generally
utilized in its formation. In the Desmidiaceae (with a solitary
exception) the gametes are set free and conjugation takes place
outside the gametangia, but in the Zygnemese the gametes unite
either within one of the gametangia or within the
connecting-tube
which joins them. In these groups the zygospore after a period
of rest forms a new gametophyte, but
in the Mesocarpeas the

zygospore immediately forms a rudimentary sporophyte with one

spore (a carpospore) the latter undergoes a long rest.
;

Much has been written concerning the sexuality of the Con-

jugata?, and much of the evidence which has been brought forward
of late years indicates that sexuality of a low type does exist.
This sexuality is less marked in some Conjugates than in others,
and least of all in the Desmidiacese. The cells in a filament of
Zygnema or Spirogyra need not necessarily be considered as all of
the same sex. It is quite possible that they have no sex until just

prior to conj ugation. There is no visible change on the conversion

of an ordinary vegetative cell into a gametangium the change is ;

a physiological one which most probably takes place immediately

antecedent to conjugation, and the formation of a male or a female
gametangium may depend upon restricted local conditions. Taking
this into consideration, it is no more surprising to see both male
and female gametes produced from the cells of a single filament
than to see a filament which gives origin to gametes of one sex only.
It has been shown that a strictly filamentous condition is of no
essential importance to 'the life of the Conjugate, and also that the
functional activities of the cells of a filament are greatly increased

during conjugation, even in those cells which take no part in the

actual conjugation 1 Thus, there is no reason why the physiological
.

1
West & G. S. West,
'
Obs. on the Couj.,' Ami. Bot. xlv, 1898, pp. 30, 3d, 37.
S
\J
9
_j
116 Chlorophycece

changes which take place in a filament on the conversion of its vege-

tative cells into gametangia changes which are rendered manifest
a
by general increased activity should not be so far modified at
different parts of the same filament as to produce a differentiation
of sex. This affords an explanation of the rare cases of " cross-
conjugation" to be described later on, but at the same time there
is no doubt that it is a general rule in the Zygnemeae that the cells
of any one filament all become of the same sex.
t/

Little is known concerning the effect of temperature and

climatic conditions on the methods of reproduction of the Coiijugatse.
A high altitude, which is usually accompanied by a relatively low
temperature, certainly favours the formation of 'cysts,' and an
increase of temperature is in many cases an aid to conjugation.
With regard to the British Islands, conjugating examples of the
Zygnemacese are more frequently met with in low-lying areas than
in upland districts, and most frequently in the western or south-
western counties. Observations tend to prove that the Conjugate
in their natural habitats withstand extremes of temperature very

well, and that they are capable of adapting themselves to very-

different conditions from those under which they normally exist.
All plants of this order, whether unicellular or filamentous, are

very slimy to the touch. This is due to the gelatinous nature of

the outer layers of the cell-wall, and this character alone frequently
enables the collector to distinguish Conjugates from most other
filamentous green Alga?.
The order is subdivided into three families, of which one the
Temnogametacea? is only known from Equatorial Africa. The
two British families are :

Family Zygnemacece. Thallus filamentous, consisting of a single series

1.

of cylindrical cells, each cell possessing one or several chloroplasts of definite

form. Any cell may become a gametangium.
Family 2. Desmidiacece. Thallus unicellular cells sometimes loosely,
;

sometimes closely joined into simple filaments, in most genera constricted into
two equal and symmetrical halves. Chloroplasts one or several in each cell,
usually of definite form. Many of the plants of this family are remarkable for
their beauty of form. Any cell may become a gametangium.

Family 1. ZYGNEMACE.E.
This family of Conjugates has a world-wide distribution and
includes some of the commonest and most striking of freshwater

Alga?. The thallus is in every instance filamentous and consists of

 Zygnemacece 117

a single series of cylindrical cells forming an unbranched filament.

Rare instances of branching are known, but the branches have
been limited to short lateral outgrowths consisting only of a few
cells such outgrowths have been observed in the genera Zygnema
;

and Mougeotia Similarly, longitudinal septa of an incomplete

1
.

character have been observed in Zygnema pachydermum West, var.

confervoides West
2
Rhizoid-like organs of attachment or haptera
.

are of frequent occurrence in young plants of Spirogyra and

Mougeotia, but have not been noticed in any of the other genera
of

ZygnemaceaB. They are usually simple or branched outgrowths

near the base of the filament, but in Spirogyra they may arise by
the modification of a conjugating-tube which has been protruded
by a cell some distance removed from those cells actually engaged

in conjugation. This is yet another proof of the increased activity

of the- filament as a whole during conjugation.
The Zygnemaceas is divided into three sub-families, of which
the Pyxisporese is only known from tropical Africa. The two
British sub-families are :

Sub-family 1. Mesocarpece. Conjugation forming a zygospore which

immediately develops a sporocarp of several cells, one of which is the
spore (carpospore). The gametophyte is developed from, this spore
after a period of rest.

Sub-family 2. Zygnemece. Conjugation producing a zygospore which

after a period of rest develops directly into a new gametophyte.

Sub-family I. MESOCARPE.E.
of this sub-family are the narrowest and most
'

The plants
delicate of the filamentous Zygnemaceae. There is a great varia-
bility in the diameter and relative length of the cells, and the
cell-wall is relatively thin.The cells contain a single chloroplast,
generally in the form of a thin axile plate, which may extend from
end to end or only occupy the median portion of the cell. Each
in a single longi-
chloroplast contains several pyrenoids arranged
tudinal series. The chloroplasts of adjacent cells usually lie in the
same plane, so that a whole filament of cells may exhibit the full
breadth of the chloroplasts or it may be in a position such that
only the thin edge of the chloroplasts can be seen. The action of
of Mougeotia.
light causes a rotation of the plate-like chloroplasts
1
West & G. S. West in Ann. Bot. 1898, xlv, p. 32, t. iv, f. 17, 18, 19 and 41.
2
West,
'
Alg. from the W. Indies,' Journ. Linn. Soc. Bot. xxx, t. xiv, f. 5.
118 CJi lo )
-

In diffused daylight they place themselves at right angles to the

direction of the incident rays, but in strong sunlight the edge of
the plate is directed towards the light. This has long been known
to students of Alga3 and special mention of it was made by Bennett.
Quite recently this phenomenon, which is well exhibited by the
chloroplasts of many green plants,
has been further investigated
and it has been shown that the chloroplast occupies on an average
Apart from the lining
1
about 30 minutes in rotating through 90 .

layer of protoplasm and the chloroplast,

a considerable proportion
of the cell-cavity is usually occupied by large fluid vacuoles.
the dissociation
Vegetative multiplication frequently occurs by
of a filament into its constituent cells, each of which forms a new

plant by rapid cell-division.

Asexual reproduction takes place occasionally in Mougeotia by
the formation of spores resembling aplanospores
2
These spores .

are produced by the division of a vegetative cell and they may be

of three
regarded as carpospores formed from sporocarps (consisting
cells) produced without conjugation, but possibly in consequence
of the stimulus which has already caused conjugation to take place
in a distant part of the filament. In the genus Gonatonema repro-
duction is solely by the formation of aplanospores, the whole of the

contents of a single cell being generally, but not always, utilized

in the formation of a spore. It has been noticed in the three best
known species of Gonatonema (viz. G. ventricosum Wittr., G. Bood-
lei W. & G. S. West, and G. tropicum W. & G. S. West) that

during the formation of the spore, and just before the rounding
off

of the protoplasmic mass, there is sometimes a more or less

3
I have care-
complete division of the cell-contents into two parts
.

studied the formation of in this genus and this

fully aplanospores
curious separation of the cell-contents into rudimentary gametes
is by no means of frequent occurrence. It is most likely a slight
retention of the last traces of ancestral sexual characters, the
of conjugation.
spores having arisen at one time by a process
Sexual reproduction of a low type takes place in Mougeotia by
conjugation. This almost always occurs between the cells of
different filaments which are lying side by side. Each cell puts
1 Lewis in Ann. Bot. xii, 1898.
F. J.
2
Wittrock, 'Om Gotl. och 01. Sotv. Alg.,' Bihang till K. Sv. Vet.-Akad. Handl.
Bd 1, no. 1, 1872, t. ii, f. 7 and 8.
s W. & G. S. West in Ann. Bot.
1898, xlv, p. 39, t. iv, f. 3; Trans. Eoy. Irish
Acad. xxxii, sect. B, 1902, p. 17, t. i, f. 5.
 ill)
Zygnemacece
out a protuberance on the side towards the other filament, and
this meets with a similar protuberance from one of the opposite
cells. The ends of the protuberances fuse together and an open
tube is formed placing the two conjugating cells in communication

with each other. This is known as the conjugating -tube. During

the development of the protuberances and their ultimate fusion
the greater part of the protoplasmic contents of each cell, including
the chloroplast, contracts away from the wall of the gametarigium
and passes into the conjugating-tube. There, a coalescence of the
gametes takes place, resulting in the formation of a zygospore,
which, although a compact mass situated in the conjugating-tube,
is not at first cut off from the rest of the protoplasm left in the
gametangia. Wittrock regards the entire H -shaped structure,

consisting of the two gametangia and the conjugating-tube,

as the

zygospore. Cell-walls soon appear in this structure, indicating the

formation of a rudimentary sporocarp, and they cut off a central
or
fertilecarpospore from the surrounding sterile cells (two, three,
four in number). Thus, the Mesocarpea? afford an instance in the
'

green Alga? of a sporophyte generation and a rudimentary

alterna-
tion of generations.'
The method of spore-formation met with in Mougeotia differs
from that which occurs in the Zygnemese. The spore is
materially
not formed by the development of a new cell-Avail around the fused
walls which cut off
gametes, but by the appearance of partition
some part of the H -shaped structure in which fusion of the gametes
has taken place. Thus, the spore in the Mesocarpeas can be re-
garded as a carpospore which is bounded partly by
new walls and
the walls of the gametangia or the conjugating-tube.
partly by
The type of conjugation between the cells of distinct filaments
is known as scalariform conjugation. In some cases conjugation
occurs between adjacent cells of the same filament. This type is
known as lateral conjugation, and although commonly met with in
the Zygnemea?, very rarely observed in the Mesocarpea^.
is

Irregularities are sometimes met with in the conjugation of

Mougeotia. Cases have been observed in which the terminal cell
of a filament has entered into conjugation through it free end, no

conjugating-tube being developed, and rare instances occur in

which three cells, each belonging to distinct filaments, have entered
in which
into conjugation. Equally rare are the hybrid examples
has occurred between species of Mougeotia of different
conjugation
120 Chlorophycece
thickness. Two carpospores have been observed to be formed in
one sporocarp of Mougeotia cajmcina', this is precisely analogous
to the double spores met with in certain Desmids and in abnormal
cases of Spirogyra.

Fig. 44. A, Mougeotia sp., from Frizinghall, W. Yorks., young filament showing
organ of attachment ( x 100). B, Mougeotia capucina (Bory) Ag., from the
New Forest, Hants., showing edge of plate-like chloroplast ( x 430). C, M.
viridis (Kiitz.) Wittr. ( x445). D H, M. parvula Hass.; D F, from Eppiug
Forest, Essex ;
G and H, from near Settle, W. Yorks. ( x 445). I, M. gracil-
lima (Hass.) Wittr., from Esher West-end Common, Surrey ( x 445). cp, carpo-
spore sp, sporocarp; z, zygospore (of Wittrock).
;
 Zygnemaeece 121

Indications of sexuality are to be found in the Mesocarpese,

but they are much less marked than in the Zygnemege. The
spores are often seen to be situated nearer to one gametangium
and the conjugating- tube of that gametangium to be thicker and
shorter than that of the other; hence the formermay be looked
upon as a female cell and the latter as a male cell. As these
indications of sexuality are scarcely discernible and often absent,
the Mesocarpea? may be regarded as having lost almost all traces
of differentiation of sex.
Genus Mougeotia Ag., 1824. [Staurospermum Kiitz., 1843 ;

Mesocarpus Hass., 1845 ;

Oraterospermum Braun, 1855 Plagio- ;

spermum Cleve, 1868.] The thallus consists of cylindrical tin-

branched filaments of elongated cells. The single chloroplast is
disposed as an axile plate, extending from end to end of the cell or
only occupying the median portion. The pyrenoids are numerous
and usually arranged in a single series. In M. capucina the
chloroplast sometimes assumes the form of an irregular axile rod,
connected with the lining layer of protoplasm by fine colourless
strands, and the vacuoles contain a purple cell-sap. In some
species the carpospores are spherical, but in others they are quad-
rate and more or less flattened, with rounded or truncated
angles.
Species of this genus were at one time referred to various genera,
such as Mesocarpus, Staurospermum, etc., according to the disposi-
tion of the sterile cells of the
sporocarp and the form of the carpo-
spores,but all the supposed generic differences have been found
by Wittrock to be present in one and the same filament of

Mougeotia calcarea Wittr. Throughout the entire genus there is

great variability in the relative size of the carpospore and the
sterile cells of the sporocarp.
In mountain tarns and lakes species of this genus are extremely
abundant, and they flourish in the summer months in small pools
on the mountains up to 3,000 ft. elevation. In these situations
the plants rarely conjugate and they are kept alive through the
'
winter largely by the formation of resting-cells or cysts,' which
are of the same form as the ordinary vegetative cells. In the
plankton of large lakes the filaments are often much twisted and
coiled.

There are about 15 British species pf the genus, of which M. scalaris Hass.
(diameter of fil. 32 35 /*) is the largest and M. elegantula Wittr. (diameter of
fil. 3'5 4'5 p.) is one of the smallest. The two most abundant species are
122 Chlorophycece
M. parvula Haws. (fig. 44 D H) and M. gracillima (Hass.) Wittr. (fig. 44 I),
which conjugate freely in all parts of the country and at all elevations up to
is a frequent species in ditches and ponds.
1,200 ft. M. genuflexa (Dillw.) Ag.
M. gelatinosa Wittr. is one of the rarest British species, being characterized
investment of the carpospore.
by the curious gelatinous
Genus Gonatonema Wittr., 1878. The thallus is similar in

all respects to that of Mougeotia, but the reproduction

is brought

about solely by the formation of aplanospores, the filaments usually

at the points of location of the spores. Each
becoming genuflexed

Fig. 45. A-F, Gonatonema Boodlei W. & G. S. West, from Mitcham Common,
Surrey. G J,G. ventricosum Wittr., from the river above Crolly Bridge,
Donegal, Ireland. (All x 445.) a, aplanospore.
 Zygmmacece 1 23

spore is formed by the rejuvenescence of the contents of the

mother-cell, and there is a greater difference in size between the
spores of species of this genus than can be accounted for by the
difference in cubical capacity of the mother-cells.

Plants of this genus are of much rarer occurrence than those of Mougeotia ;

in fact, they are amongst the rarest of the Conjugatee. G. notdbile (Hass.)
Wittr., with vegetative cells 12 15 yn in diameter, has not been seen since its
discovery by Hassal in 1845. G. vcntricosum Wittr. (fig. 45 J), with veg. G
-
cells 7 5 8'6 fj.
in diameter, is known from Ireland ;
and G. Boodlei W. &
G. S. West (fig. 45 A
F), with veg. cells 5 5 '5 /* in diameter, has been found
in Surrey. The aplanospores of G. ventricosum are likewise considerably
different from those of G. Boodlei and they are sometimes produced by the

swelling of the free end of the terminal cell of a filament.

Sub-family II. ZYGNEME/E.

The plants of this sub-family consist of unbranched filaments
of cells similar to those of the Mesocarpese, but usually of much

larger size. There is a lining layer of protoplasm in each cell and

the nucleus is situated in the central portion of the cell, embedded
in a mass of protoplasm which is connected with the lining layer
by numerous radiating strands. Gerassimoff finds certain cells of J

Spirogyra majuscula to contain two ordinary nuclei or one com-

pound nucleus. The chloroplasts, which contain prominent pyre-
noids, are somewhat variable in form and disposition in Debarya ;

there is a single axile plate similar to that of Mougeotia; in

Zygnema there are two star-shaped chloroplasts suspended in the
middle line of the cell and in Spirogyra the chlorophyll is
;

arranged in one or more bands twisted spirally round the interior

of the cell-wall. The chloroplasts of Spirogyra are very variable
and special reference is made to this feature under the genus.
Vegetative multiplication takes place as in the Mesocarpeae by
the breaking up of a filament into its constituent cells, or into
groups of cells, which then grow into new filaments.
Asexual reproduction sometimes takes place by the formation
of aplanospores, which are produced from the contents of a single
cell.
They are more frequently found in Zygnema than in Spiro-
gyra, and until the discovery of Z. spontaneum Nordst. in West
Africa 3 the only known method of reproduction of this species was
,

1
Gerassirnoff in Bull. Soc. Imp. Nat. Moscou, 1897.
-
W. & G. S. West in Journ. But. Febr. 1897, p. 15.
1 24 ChlorophycecK
an asexual one The aplanospores (or, as they are frequently
1
.

termed, parthenospores ') of Zygnema are generally globular and

'

rather smaller than the zygospores, but those of Spirogyra are as

a rule very similar in form to the zygospores. Klebs has stated
that parthenogenetic resting-spores can be produced in filaments
with long conjugating- tubes by placing them in a strong solution
of sugar. Spirogyra mirabilis (Hass.) Petit is reproduced solely
by spores resembling aplanospores, but produced by a degenerate
form of conjugation'-. Many of the upland forms of Zygnema
' '

habitually form solitary resting-cells or cysts in order to survive

the winter.
Sexual reproduction takes place by conjugation in all the

genera of this sub-family. The usual type of conjugation is scala-

riform, between the cells of two distinct filaments. The conju-

is formed exactly as in Mougeotia, and in all except
gating-tube
species of the genus Debarya and certain species of Zygnema
belonging to the section Zygogonium, the fusion of the gametes
takes place in one of the gametangia. In the formation of a
gamete the protoplasmic mass contracts away from the cell-wall,
the chloroplast loses much of its original form, and the mass
becomes of an ellipsoidal shape. The whole of this mass then
glides gradually from its gametangium through the conjugating-
tube into the opposite gametangium. The gametes may coalesce
immediately on contact and before the one has completely passed
through the conjugating-tube, or they may lie side by side in the
gametangium before fusing. On the coalescence of the gametes
and the fusion of their nuclei the mass generally assumes a spheri-
cal or ellipsoidal form,develops a thick cell-wall and is known as
the zygospore (or zygote).
The cell in which the fusion of the gametes takes place and
therefore the one in which the zygospore is formed, is known as
the female cell, and the one which is emptied as the male cell ;

and that part of the conjugating-tube developed from the female

1
Nordst. 'Alg. et Char. Sandvic.' 1878, p. 17, t. 1, f. 23, 24.
-
Petit in his Hpirogyra des envir. de Paris,' p. 14, writes concerning S. mira-
'

bilis "Cette tres curieuse espece ne conjugue pas et ne laisse voir aucun tube
:

copulateur; a une certaine epoque de la vie de la plante, les cellules renflent vers le
milieu, 1'endochrome se partage en deux parties qui se concentrent sous forme de
globule aux deux extremities de la cellule il se forme ainsi une differentiation entre
;

les parties de 1'endochrome. Bientot les deux globules se rapprochent vers la partie
reuflee de la cellule et finissent par se reunir en constituent ainsi la zygospore."
This is a similar phenomenon to that which occasionally occurs in the formation of
the aplanospores of Gonatonema. (Consult page 118.)
 Zygnemacece 125

cell isusually thicker and shorter than the part developed from
the male cell. The female cells frequently become much swollen
after the fusion of the gametes.
On the examination of a large number of conjugated examples
of either Spirogyra or Zyynenta one feature must impress itself
even on a casual observer, namely, that "the direction of conjuga-
tion is clearly governed by some physiological law, the movement

of protoplasm between the two filaments almost invariably taking

place in one direction only, so that one of the two conjugating

filaments is entirely emptied, while the other is filled with zygo-
Thus, the cells of one filament usually become all of
1
."
sperms
the same sex.
Two filaments are generally concerned in an example of scala-
riform conjugation, but three, four, five, six, or more, are occasion-

ally seen (vide fig. 49 A). Such are mostly cases of polygamy or
polyandry, and statistics seem to show that the former is the more
frequent.
Lateral conjugation, or conjugation between adjoining cells of
the same filament, is frequently observed, but it is much scarcer
than scalariform conjugation and may be regarded as exceptional.
It more often met with in Spirogyra than
is in Zygnema and is

especially frequent in some of the smaller species, such as 8.

tenuissima (Hass.) Kiitz., S. inflata (Vauch.) Rabenh., and S.
'

varians (Hass.) Klitz. Cross-conjugation, in which some cells of

the filaments are male and others female, and therefore in which
perfectly normal zygospores are found in both filaments, is exceed-
ingly rare. In both lateral and cross-conjugation, as was mentioned
in the general remarks under the order Conjugate, a differentia-
tion of sex must take place amongst the cells of the same filament,
some becoming male and others female.
Sometimes certain irregularities are met with in the conjuga-
tion, and these are no doubt largely due to the increased functional
is such a striking
activity of all the cells of the filaments which
accompaniment to conjugation. Zygospores are occasionally ob-
served which have been produced by the coalescence of three
2
gametes, two male and one female but attempts of this nature
,

1
Bennett and Murray, 'A Handbook of Cryptogamie Botany,' London, 1889,
p. 266.
-
West in Notarisia 1891, vi, t. siii, f. 13 ; Borge in Bihang till K. Sv. Vet.-
Akad. Handl. Bd 17, 1891, no. 2, t. i, f. 2; W. & G. S. West in Ann. Bot. xii,
1898, t. v, f. 66.
 126 CMorophycece
are usually abortive 1 (fig. 49 B). Gerassimoff has observed the
conjugation of two female cells with one male cell and the forma-
by the coalescence of -the protoplasm of the
tion of the zygospore
male with that of one of the female cells, a parthenospore
cell

being formed in the other female cell. In those plants of the

Zygneme* in which the zygospore is formed in the conjugating-
tube, conjugation between three cells entails the production of
two zygospores, each of which is less than the normal size. Owing
to sudden changes of local conditions it frequently happens that
the conjugation has been brought to an abrupt termination before
the proper formation of the zygospores has taken place. In such
cases of interrupted conjugation the spores are apt to be very
variable sometimes the spore is not of its true form, and at other
;

times one small one is formed in each gametangium or two small

ones in the female gametangium 2 The discovery by Gerassimoff .

of binucleated cells in the Zygnemeae may perhaps afford an expla-

nation of some of these irrregularities of conjugation. He states
that in the conjugation of binucleated cells parthenospores were
sometimes observed.
Rare instances of hybrids are known, in which conjugation,
with the production of zygospores, has occurred between filaments
of different species 3 .

The
ripe zygospores possess a thick wall of cellulose which is
divisible into three coats, the outer one being cuticularized and
the middle one usually exhibiting some form of sculptured mark-
ings. The contents of the zygospore often turn red in colour and
develop a large amount of oily material. Sometimes the chloro-
in the zygospore some-
plasts of the two gametes remain intact ;

times those of the female remain intact and those of the male
disintegrate but it is quite a common thing for the chloroplasts
;

to completely disintegrate before the fusion of the gametes. The

zygospores undergo a period of rest, surviving the winter or even
a period of drought, and they usually germinate after the conju-

1
W. & G. S. West, 1. c. figs. 67 and 69 Schmula in Hedwigia, xxxviii, 1899.
;

(c. fig.) Copeland states that in such a case the nucleus of the abortive male cell
is situated against the wall opposite, remote from the conjngating-tube. (Cf. Bull.
Torr. Bot. Club, xxix, 1902.)
2
Eosenvinge in Ofvers. K. Vet.-Akad. Forh. 1883, no. 8, t. viii, f. 111; West
in Notarisia 1891, vi, t. xiii, f. 27, 28; Hansgirg in Hedwigia 1888, t. x, f. 6;
W. & G. S. West, 1. c. t. v, f. 7480.
3
Cf. Spirogi/ra maxima var. iiHcqudliis Wolle Freshw. Alg. u. s. t. cxxxviii,
f. 5, 6; t. cxlii, f. 5, 6; W. & G. S. West, 1. c. t. v, f. 70, 71.
 Zygneniace' 127

gating filaments have entirely perished. In Spirogyra the two

outer coats are ruptured, generally at one end of the spore, and
the protoplasm is protruded as a filament clothed with the inner-
most cellulose wall. This filament becomes divided by a transverse
septum, one cell becoming an organ of attachment and containing
little or no chlorophyll, whereas the other cell increases in size,

contains one or more chloroplasts, and by repeated divisions forms

a filament. The distinction between base and apex is soon lost
and the filament floats freely in the water.
Organs of attachment
have, however, been observed to be developed subsequently from
older cells. In the germination of zygospores of Zygnema the
distinction into base and apex is
scarcely evident.
Certain of the zygospores of Spirogyra velata Nordst. have
been observed to germinate immediately after their formation
without the development of a thick cell- wall (fig. 49 E and F) 1
.

Genus Debarya Wittr., 1872.

[Mougeotiopsis 1894.] Palla
The thallus consists of simple, cylindrical filaments, with or without
a distinct constriction between the cells, and with a thin external
mucous covering. There is one chloroplast in each cell, disposed
as an axile plate similar to that of Mougeotia, and containing
several pyrenoids. In rare instances pyrenoids are absent. The
gametes are formed from the entire contents of the gametangium
and conjugation takes place in the conjugating- tube, the mature
zygospore occupying a position between the gametangia. The
zygospores are very variable in character and there are consider-
able differences in the actual processes of conjugation, but in one
curious fact
all the
species agree. A
peculiar change comes over
the empty gametangia as the zygospore is being formed;
they
become very clear and refractive, and sometimes a series of stria-
tions become visible parallel to the transverse walls. have They
at this stage the appearance of
solidity, most likely owing to the
deposition of annular thickenings of cellulose inside the cell-wall
on the receding of the protoplasm during This
conjugation.
feature is noticeable not only in
living specimens, but also in old,
preserved examples.

All the species of this genus are of rare occurrence. D. glyptosperma (De
Bary) Wittr. is more widely distributed than the others; the cells are 9 15 ^
in thickness and 6 15 times longer than the diameter the tubes ;
conjugating-

1
W. & G. S. West iu Ann. Bot. xii, 1898, t. v, f. 84, 85.
128 Chlorophycece
are often very long and the zygospore is large and ellipsoidal, frequently with
one or two mamillate protuberances at the poles length of zygosp. 35 72 /*,
;

breadth 1640 ^ (fig. 46 A). D. Icevis (Kiitz.) W. & G. S. West is a larger

species with shorter cells, which are 20 26 ju
in thickness and 2| 4 times

Fig. 46. A, Debari/a gli/ptox-perma (De Bary) Wittr., from Fairfield, Westmoreland
(x275). B, D. calospora (Palla) W. & G. S. West, from Pilmoor, N. York*.
( x430). C E, D. hci-is (Kiitz.) W. & G. S. West, from Mitchain Common,
Surrey; C, x 180; D, x250; E, mature zygospore, x 430. F I, D. Desmidi-
oides W. & G. S. West, from near the Lizard, Cornwall ( x 430).

longer than the diameter. The zypospore is ellipsoidal (44 50 p x 29 36 p.)

with a scrobiculate middle coat (fig. 46 C E). D. calospora (Palla) W. &
G. S. West is 11 13 p. in diameter; the zygospores are subglobose or ellip-
soidal, 18 26 /x in diameter, and ornamented with large scrobiculations
 ZygnemaceoB 129

(fig.46 B). Palla 1 placed this species under a new genus of the Conjugate
owing to the absence of pyrenoids in the plants he observed. More recently,
however, precisely the same species has been discovered ivith pyrenoids, and
as the latter are subject to much variability their presence or absence is of no
2
generic value The most remarkable species of the genus is D. Desmidioides
.

W. & G. S. West 3 which, up to the present time, has only been observed
,

from Cornwall. The cells are short, only 2^ 65 times longer than the
diameter and 7 7 8-6 /* in thickness (fig. 40 F I). There is a most evident
-

constriction between all the cells of the filaments, and the latter dissociate
with the greatest ease into their individual cells, conjugation only occurring
between the free, dissociated cells. This plant throws much light on the
origin of the Desmidiacese.

Genus Zygnema Ag., 1824. [Zyyogonium Kiitz. 1843.] The

filaments are simple, consisting of a single series of cylindrical
cells placed end to end, and sometimes exhibiting a slight con-

striction at the points of junction. Each filament possesses an

external mucous coat which is sometimes remarkable for its size
and strength. There are two star-shaped chloropla.sts suspended
in the median line of each cell, each one containing a single

large pyrenoid. Sometimes the chloroplasts are very indefinite,

their form and disposition being difficult to make out. This is
particularly the case in Z. ericetorum (Kiitz.) Hansg., some forms
of which greatly resemble species of Microspora or of Rhizoclonium.
The coalescence of the gametes takes place either in one of the
gametangia (the female) or in the conjugating-tube. Species in
which the latter occurs were referred by Kiitzing to a separate
genus Zyyogonium. The zygospores are globose or ellipsoid.

There are about a dozen British species of this genus, of which Z. ericetorum
(Kiitz.) Hansg. (fig. 47 C) is the most widely distributed. This species lives
equally well in water or on damp heaths or peaty moors, and it frequently
assumes a purple or violet colour owing to the formation of phycoporphyrin
in the cell-sap. It is an Alga which fulfils an important function on some of
the heaths and moors. In the drier and hotter periods of the year, thickly-
matted sheets of Z. ericetorum, often many square feet in extent, are found
covering wide patches of almost bare sand or peat, round such plants as
Drosera, Carices, etc. These mats of Zygnema have great absorptive capacity,
greedily taking up water, and in this way they regulate the moisture of the
surface soil, the thriving of some of the smaller Phanerogams depending to a

1
E. Palla, Ueber eiue neue, pyrenoidlose Art und Gattimg der Conjugate!!,
' '

Berichte Deutsch. Bot. Gesellsck. xii, 1894, Heft 8, pp. 228236, t. xviii.
-
W. & G. S. West in Aim. Bot. xii, 1898, p. 49 ; in Journ. Bot. Aug. 1900,
p. 289.
3
W. & G. S. West iii Journ. Bot. 1903, p. 7 (Sep.), t. 446, f. 19.
W. A. 9
130 CMorophycece
1.
great extent on the presence of the Zygnema Z. ericetorum very rarely con-

jugates, and mature zygospores, which are found in the conjugating- tube, have
only been observed on one or two occasions. The filaments are 1"> 22 p in
diameter and the cells often become irregularly thickened.

A, Zygnema st ell in urn (Vauch.) Ag., from Cam Fell, W. Yorks.

x 430).
Fig. 47. (

B, Z. Vaucherii Ag. var. stagnate (Hass.) Kirchn., from near the Lizard,
Cornwall (x430). C, Z. ericetorum (Kiitz.) Hansg., from Bombald's Moor,
W. Yorks. (x415). D, Z. leioxperninm De Bar.y, from Esher Common, Surrey
(x430), a portion of a filament which was conjugating along the greater part
of its length. E, Z. insigne (Hass.) Kiitz., from Malham, W. Yorks. ( x 330).
F, Z. Ealfsii (Hass.) De Bary, from Chippenham Fen, Cambridgeshire ( x 430).
ap, aplanospore.

1
This phenomenon is much more evident in some parts of the tropics, and
was by Welwitsch in the Journal of Travel and Natural
'
attention first called to it

History,' vol. i, 1868. In the damp sandy valley of the Cuanza Kiver, in Angola,
the Alga Porphyrosiphon Notarisii occurs in extensive sheets, closely spread like a
 Zi/gnemacece 131

Z. pectinatum (Vauch.) Ag., Z. cruciatum (Vauch.) Ag. and Z. insigne

(Hass.) Kiitz. (fig. 47 E) are the three largest British species, usually occurring
in ponds or in road-side ditches. The smallest form of the genus is Z. Vauch-
erii Ag. var. stagnate (Hass.) Kirchn. (fig. 47 B), the filaments of which reach
a maximum thickness of 13 /u. Z. anomalum (Hass.) Cooke is a species
largely confined to upland bogs and is remarkable for its wide mucous in-

vestment.

Genus Spirogyra Link, 1820. This is the best known and

most abundant genus of Conjugate, and it has a distinct preference
for low-lying, quiet waters,
such as those of ponds
and ditches. The fila-
ments are quite simple
and occur in bright green
flocculent masses, often
several feet in diameter.
The cells are cylindrical
and they exhibit great
variability both with re-
gard to their diameter
and their relative length.
The cell-wall is firm, with
an outer mucous coat
which renders the plants
very slimy. In most
species the transverse
cell-walls are quite plane,
but in some of the smaller
species there is a curious
Fig. 48. A, Spirogyra majuscula Kiitz., from
annular ingrowth of cel- Coates, Gloucestershire, single cell showing chloro-
plasts and nucleus ( x 300). B, Spirogyra sp.,
lulose at the ends of each from Coates, Gloucestershire ( x 370). C, S. tenuis-
cell(wdefig.48C). When
sima (Hass.) Kiitz., from Mitcham Common, Surrey
(
x 390). ,
nucleus.
this is present the cells are
"
said to possess replicate ends." It is a character which is constant
for the species in which it is found, although the ingrowths are not

net over the intergrown with small herbaceous plants and shrubs. The mat-
soil,
like sheets of theAlga eagerly absorb the atmospheric moisture during dewy nights,
affording by this means a refreshing protection to the roots of many other and
larger plants during the glowing heat of the following day. Welwitsch states that
the growth and thriving of the numerous small phanerogamous plants in these
places is conditional on the co-presence of the Alga. Cfr \V. & G. S. West in
Jouru. Bot. 1897, p. 303.

92
132 Chlorophycece

necessarily present at the extremities of every cell in the filament.

The nucleus is situated in the central portion of the cell and the
protoplasm in which it is embedded is connected with the lining
primordial utricle by numerous delicate strands. Many of these
strands reach the lining protoplasm exactly opposite a pyrenoid,
and as the starch formed during carbon-assimilation makes its

appearance round the pyrenoids, this fact has been brought forward
as a reason for supposing the nucleus to have a direct function in
starch-formation. The nucleus is often clearly visible in living
specimens, particularly of 8. majuscula Klitz. and 8. pellucida
(Hass.) Klitz. Mitotic division has been observed in this nucleus
by Mitzkewitsch and by C. van Wisselingh, and Gerassimoff has
observed cells with either a compound nucleus or two ordinary
nuclei.
The chloroplasts are the most striking feature of Algge belonging
to this genus. They are disposed in the primordial utricle in the
form of spiral bands, and they vary in number from one to about
six or seven in each cell. In some species they are coiled into
very close spirals, but in others they are practically straight and
longitudinal in some, as in S. neglecta (Hass.) Kiitz., their margins
;

are quite smooth and there is a regular axile series of pyrenoids ;

in others, as in S. nitida (Dillw.) Link or S. porticalis (Vauch.)

Cleve, the spiral bands are very broad, with serrated margins and
scattered pyrenoids. Between these two extremes there is every
gradation, and the character of the chloroplasts always remains
constant for any particular species, even though the number of
them may vary in different cells of the same filament. It has been
ascertained by Kolkwitz 1 that the chloroplasts grow in length in
the direction of the coils by both apical and intercalary growth;
and therefore, as this is obliquely to the surface of the cell-wall,
there is a gliding motion of the spiral bands through the primordial
utricle.
The coalescence of the gametes takes place in all cases in the
female gametangium, which often swells up to twice its normal
size. The zygospores may be globular, ellipsoidal, cylindrical with
obtuse ends, or they may more rarely assume the form of
flattened discs ;
and the middle coat of the zygospore is frequently
ornamented with distinctive markings.

1
Kolkwitz in Festschr. f. Swendener, 1899, pp. 271287.
 Zygnemacece 133

: !

Fig. 49. A, Spiroriyra nitida (Dillw.) Link, from near Morecambe, Lancashire; scala-
riforra conjugation between six filaments ( x 75). B, S. setifonnis (Eoth) Kiitz.,
showing frustrated attempts at the conjugation of two male cells and one
female cell ( x 90). C, S. Spreeiana Babenh. from near Esher, Surrey ( x 390).
,

D, S. iiiflata (Vauch.) Rabenh., from near Esher, Surrey; showing lateral

conjugation x 390). E and F, zygospores of S. velata Nordst., from Shipley
(

Glen, W. Yorks., germinating immediately after their formation and before the
differentiation of the wall of the zygospore ( x 165). G, germination of zygo-
spore of S. velata after normal period of rest (
x 230).
134 ChlorophycecK
There are about 24 British species of the genus, S. gracilis (Hass.) Kiitz.
being the smallest, with a minimum thickness of about 10 /i, and S. crassa
Kiitz. the largest, with a maximum thickness of about 150 /j. Several species,
such as S. tenuissima (Hass.) Kiitz. (fig. 48 C), S. gracilis (Hass.) Kiitz.,
S. communis (Hass.) Kiitz., S. varians (Hass.) Kiitz. and S. nitida (Dillw. !

Link (fig. 49 A), are abundant in every part of the British Islands except in
the most mountainous districts. S. velata Nordst. and S. calospora Cleve are
notable for their beautifully marked zygospores.

Plants of this genus often cause trouble in cress-beds, the

matted masses of Spirogyra preventing the growth of the cress
plants.
Genus Choaspis S. F. Gray, 1821. [Sirogonium Kiitz., 1843.]
It is very doubtful if this genus should be separated from Spirogyra.
The filaments are similar
to those of many species
of the latter genus and
the chloroplasts, although
more or less straight
and longitudinal, are not
straighter than those of
Spirogyra majuscula or
S. pellucida. There are,

however, certain constant

distinctions. There is a
remarkable absence of
the external mucous coat,
in consequence of which
this is the only Conjugate

Fig. 50. A C, Choaspi* xtictica (Eng. Bot.) which is not slimy or

O.K., from Ingleton, W. Yorks. (x 120). The process of
slippery.
the female
conjugation is peculiar. The gametangia, more especially
one, become swollen and sometimes genuflexed. The conjugating-
tubes are not distinct, but the walls of the gametangia come into
apposition and a perforation
formed at the point of contact.
is

The chloroplasts also completely disintegrate, even before the

The
gametes have receded from the walls of the gametangia.
coalescence of the gametes takes place in the female gametangium.
a few of the
Conjugation only appears to affect indiscriminately
cells of the filament, and as these cells are usually shorter than the

ordinary vegetative cells, they may perhaps

be specially set apart
as sexual cells, almost comparable to those which are specially cut
 Desmididcece 135
1
off in the TemnogametaceEe. Gray's description of Choaspis is
a very good one and is twenty-two years previous to Klitzing's
description of Sirogonium.
Ch. stictica (Eng. Bot.) O. Kunze [ = Sirogonium sticticum Kiitz.] is the only
British species (fig. 50 A C). The cells are 2 6 times longer than their
diameter (which is 40 50 /*), and there are several more or less longitudinal
chloroplasts. The zygospores are ellipsoidal, about 75 /* in length and 42 p. in
breadth. The plants usually occur attached to stones over which the water
is running fairly fast ;
they are also said to occur in stagnant water.

Family 2. DESMIDIACE.EJ.
Theplants included in this family are remarkable for their
great diversity of form and their wonderful symmetry. Indeed,
the group includes some of the most beautiful of microscopic
objects. They are unicellular plants and the major portion of
them lead a solitary existence. Certain of them are, however,
associated in colonies and others are more or less closely united
into long filaments. They are essentially free-floating Algae and
frequently occur in enormous abundance in small ponds, in the
quiet margins of lakes, and in other favourable localities.
Most Desmids exhibit a more or less distinct constriction into
two perfectly symmetrical halves each half is termed a semicell
;

and the narrower part connecting the two semicells is known as

the isthmus. The angle resulting on either side from the con-
striction or narrowing of the cell is known as the sinus.
One of the most striking features of the family is the extra-
ordinary complexity of the cell-outlines. The cell is often deeply
lobed or incised, and the exterior of the cell-wall is frequently
covered with granules, spines, wart-like thickenings, or other pro-
tuberances, most of which have a definite pattern of arrangement.
This firmer portion of the cell-wall consists chiefly of cellulose, and
external to it are layers of gelatinous pectose compounds. The
latter often form a thick mucilaginous coat completely surrounding
the individual, or, as in the case of colonial forms, entirely envelop-
ing the colony. It is by means of this mucilaginous envelope
that Desmids adhere to other larger aquatic plants, -and sometimes,
when the conditions have been favourable for rapid multiplication,
enormous numbers of individuals occur embedded in masses of
jelly.Sometimes the mucus is very tough. In the genus Spon-
1
S. F. Gray, Arrang. Brit. Plants, 1821, vol. i, p. 299.
136 Chlorophycece

dylosium the cells are united into filaments by mucous threads

passing between their apposed ends, and the filaments break much
more readily across the isthmus of a cell than at the points of
apical attachment.
The minute structure of the cell-wall was first studied by
Klebs who examined the nature of the gelatinous outer coat and
1

also demonstrated the presence of pores in the cell-wall. Shortly

afterwards Ltitkemuller described the pores in the cell-wall of
2

Closterium and quite recently he has published a very important

account of the structure of the cell-wall of Desmids
3
.

In a few Desmids those belonging to the Spirotsenieae the

cell-wall consists of a single layer of cellulose of homogeneous
structure, but in the majority of Desmids it composed of two
is

well differentiated layers. The inner layer is structureless and

consists of cellulose ;
the outer layer is stronger and thicker, con-

sisting of a ground
substance of cellulose through which pass
numerous tube-like structures or 'pore-organs.' The latter are
not of cellulose, and a small pore or canal runs through
composed
the middle of each one. The contents of these canals also traverse
the inner layer of the cell-wall and often terminate on its inner
surface in lens-shaped or bulbous swellings. From the outer end
of the pore-organs there often arises a delicate flower-like or club-

structure through which the canal passes. More often this

shaped
structure is entirely replaced by a small perforated
wanting or is

button or rod. In many of the larger Desmids there are numerous,

smaller, thread-like pores between the larger
canals and only
Outside the differen-
traversing the outer layer of the
cell-wall.

tiated parts of the cell-wall is the mass of gelatinous pectose

compounds present in so many species, and which

is secreted by

the protoplasm and passed outwards through the pores. This

outer gelatinous coat often exhibits a prismatic or fibrillar struc-
ture and is frequently the home of epiphytes or of numerous rod-

shaped bacteria. The cell-wall sometimes contains a trace of

silica.

With the exception of the lining primordial utricle the

Biol. Ceutralbl.
1
Klebs, 'Ueber Bewegung und Schleimbildung der Desmidiaceen,'
Bd v, 1885.
Oesteir.
Lutkemiiller, 'Die Poren der Desmidiaceen Gatt. Closterium Nitzsch,'
2

botan. Zeitschr. Bd 44, 1894.

Lutkemiiller, 'Die Zellmembran der Desmidiaceen,' Beitrage
;i zur Biol. der
Pflanzen, herausgegeben von F. Cohn, Bd viii, Breslau, 1902.
 Desmidiacece 137

cytoplasm is variable in its general disposition, the variability

depending largely on the nature and arrangement of the chloro-
plasts. In those Desmids with large, central chloroplasts, vactioles
may be absent or much reduced, and they are often confined to
one at each extremity of the cell in others with parietal chloro-
;

plasts large vacuoles are often present in the central portions of the
cell. The general transparency of the cell- wall enables the circula-
tion of the protoplasm to be seen extremely well, especially in the

larger species. The nucleus is usually embedded in a small mass

of protoplasm in the median part of the cell in the region of the
isthmus, and in some genera is readily visible without the use of
reagents. In the genera Gonatozygon, Closterium and Pleuro-
tcenium, and in certain species of the genus Penium, there is a
well marked terminal vacuole at each extremity of the cell, con-

taining one or many moving granules. These granules, which

may be quite irregular in form or of some definite shape, exhibit
rapid vibratory movements and have in the genus Closterium been
shown to be minute crystals of gypsum. Their movements cease
immediately on the death of the protoplasm.
If Desmids are kept living in small glass vessels for some
time, and therefore under abnormal conditions, curious changes
frequently occur in their protoplasm, resulting in the formation
of large vacuoles which previously did not exist. These vacuoles
generally contain numbers of minute moving corpuscles which
are somewhat different in appearance from those normally present
in the apical vacuoles of Closterium. As many as six large
vacuoles can be noticed in a single semicell of Pleurotcenium
coronatum (Breb.) Rabenh., each one being partially filled with
an incessantly moving mass of minute corpuscles, which move
freely in the vacuole and always collect towards its base. These
corpuscles or granules are of a pale yellow colour and appear
brown in a thin stratum but when present in immense numbers
;

they give the Desmid almost a black appearance. Under abnormal

conditions these
moving granules are developed in numerous
genera, such as
Penium, Cosmarium, Euastrum, Micrasterias,
Staurastrum, Arthrodesmus, etc., and at the same time the cell-
sap in the vacuoles often becomes coloured violet with phycopor-
phyrin, a pigment which occurs normally in the cell-sap of very
few Desmids.
The form and disposition of the chloroplasts are conspicuous
138 Chlo rophycece.

features of many Desmids. They may be situated in a central

position in the cell or semicell, or they may take the form of

parietal cushions or bands. In the cells of Spirotcenia, Mesotce-
nium, Roija, and in some forms of Gosmocladium, Gonatuzygon and
Penium, there is
only one chloroplast, and the nucleus conse-
quently occupies an asymmetrical position. The majority of
Desmids possess two centrally disposed (axile) chloroplasts, sym-
metrically arranged, one in each semicell, but in Pleurotcenium
and a few species of Cosmarium, Staurastrum and Xanthidium
there are a number of parietal, cushion-like chloroplasts in each
semicell. The chloroplasts are very variable in character and dis-
position, and many intermediate conditions are noticed between
truly axile and truly parietal masses of chlorophyll. There can
be little doubt that the axile chloroplast was the original type and
that the parietal condition has been secondarily acquired by
certain Desmids. In those forms in which the cell is deeply lobed
or incised the chloroplasts often closely follow the cell-outlines,

being themselves symmetrically lobed. Pyrenoids are present in

the chloroplasts of all Desmids. In most forms one or two pyre-
noids are present in each semicell, but in the large flattened forms
of Euastrum and Micrasterias, and in the elongated cells of
Closterium, Penium, Pleurotceniwn, Tetmemorus, etc., the chloro-

plasts usually contain many pyrenoids. In certain species, such

as Spirotcenia acuta Hilse and Penium subtile W. G. S. West, &
only a solitary pyrenoid occurs in each cell. In certain genera the
pyrenoids are commonly subject to variation in number and dis-
position but in other genera they are remarkably constant.
1
,

Under normal conditions Desmids frequently exhibit very

active movements.
Klebs described four phases of motion in
certain elongated forms, all the movements being due to an
exudation of mucilage, the nature and extent of which has been
recently demonstrated in a few species by Schroder
2
The net .

result always a movement in the direction of the strongest

is

light, the longest axis of the Desmid being placed in the direction
of the incident rays of strong light and at right
angles to those of
feeble light.

Vegetative multiplication takes place by simple cell-division,

1
Lutkenauller, 'Beobacht. iiber die Chloroph. einig. Desmid.,' Oesterr. botan.
Zeitschrift, xliii, 1893, no. 1 ; West &
G. S. West, in Ann. Bot. xii, 1898, pp. 51,
52, t. iv, f. 3036
;
G. S. West in Journ. Linn. Soc. Bot. xxxiv, 1899, pp. 399, 400."
2
Schroder in Verhandl. d. Heidelb. Naturhist.-Med. Vereins, Bd vii, 1902.
 Desmidlacece 139

each division occupying about a day in the smaller species and

several days in the larger species. The first step in this cell-fission
is an elongation of the isthmus, causing a slight separation of the
two semicells. The elongated isthmus generally becomes swollen
and soon shows signs of a constriction midway between the two
semicells. By this time the nucleus has completely divided and

Fig. 51. A, Cloaterium Ehrenlwrriii Menegh., from Glyder Fawr, N. Wales ( x ls4l.
B, Cl. Leilileiiiii Kiitz., from near March, Cambs. ( x 334). C, Micrastcriii*
oscitans Ealfs, var. mttcronata (Dixon) Wille, from Kerry, Ireland (x!84).
D, Peniitm eucurbitinum Biss., from near St Just, Cornwall ( x 43-5). E, Stuuni-
strum Kji'lmanni Wille, from 2(500 ft. on Glyder Fawr, N. Wales ( x 435).
F, Cosmariiun Klebnii Gutw. (a form), from near Ely, Cambs. ( x 435).

shortly afterwards the constriction deepens, cutting the median

portion into two young semicells, which usually remain in contact
by their apices until they are practically full-groAvn. The orna-
mentation of the cell-wall gradually makes its appearance on the
young semicells as they assume the normal size. At first they are
140 Chlorophycece

very pale in colour, but the chloroplasts are quickly developed and
pyrenoids soon make their appearance. All Desmids which undergo
normal cell-division at the region of the isthmus consist of one of
the half-cells of the mother and a newly developed half, but in
certain species of Closterium and Penium the cell may consist of

portions of several generations.

Asexual reproduction takes place very occasionally by the
formation of aplanospores. These have been seen in Spondylosium
nitens (Wall.) Arch. 1 and in Hyalotheca neglecta Racib. 2
Sexual reproduction of a degenerate type takes place by the
conjugation of two individual cells and the formation of a zygospore.
The two conjugating cells, which in the vast majority of Desmids
are sexually indistinguishable, approximate and become enveloped
in mucus by a further secretion of the gelatinous pectose con-
stituents of the cell- wall. In many Desmids the semicells of each
individual come apart at the isthmus and the entire contents of
the cell issue as a gamete, the latter having the appearance of a
protoplasmic vesicle more or less filled with an irregular mass of
chlorophyll. In some species a protuberance of variable size arises
from the isthmus of each conjugating cell, and on the fusion of the
protuberances to form a conjugating- tube, the gametes issue into
it. The union of the gametes results in a zygospore, which
develops a cell-wall with three distinct layers. The inner layer is
thin and colourless, the middle layer is brown and firm, and the
outer layer either retains a smooth surface or becomes covered
with variously arranged warts or spines. Sometimes more than
two cells have participated in the formation of a zygospore, the
latter having been formed by the union of three 3 or even four 4

gametes.
All the filamentous Desmids dissociate into their individual
cells prior to normal conjugation, with the exception of certain
species of the genus Desmidium, and the zygospores are formed
between the conjugating cells in all Desmids except Desmidium
cylindricum Grev. In this species the zygospore is formed within
the female cell as in Spirogyra and Zygnema.

1
Wallich in Ann. Mag. Nat. Hist. ser. 3, vol. v, I860, t. vii, f. 10, 11 ;
Turner
in Kongl. Sv. Vet. -Akad. Handl. Bd xxv, no. 5, t. xviii, f. 7, 8.
'
W. & G. S. West in Ann. Bot. xii, 1898, t. iv, f. 2H 27.
:i
West in Journ. Linn. Soc. Bot. xxix, t. xxiv, f. 5 W. ;
& (Jr. S. West in
Journ. Koy. Micr. Soc. 1897, t. vi, f. 5.
4
Turner, I.e. t. x, f. 16 e.
 Desmidiacece 141

It frequentlyhappens that conjugation takes place immediately

after vegetative division and before the young semicells have
arrived at maturity and for any trace of sexuality to exist under
1
,

such conditions, one must imagine that the physiological change

Fig. 52. AC, Staurastrum Dickici Ralfs; three stages in the conjugation, from
the New
Forest, Hants. ( x 356). D
and E, Peniinn didymocarpiun Lund.;
D, conjugation of four individuals just produced by division; E, completed
conjugation showing the double zygospore; from near Balallan, Lewis, Outer
Hebrides (x464). F, Closteriitm lineatum Ehrenb., showing the double zygo-
spore, from near Glenties, Donegal, Ireland ( x 100). z, zygospore.

from the vegetative to the reproductive cell occurs immediately

antecedent to conjugation. Conjugation between adjacent cells of
filamentous species (i.e. lateral conjugation) has been observed in

1
Archer in Quart. Journ. Micr. Sci. ii, p. 251 ;
W. & G. S. West in Journ.
Roy. Micr. Soc. 1896, p. 151.
142 Chlorophycece
the genera Sphcerozosma and Spondylosium, but species of these
genera conjugate normally only after dissociation of the filaments.
The four Desmids Glosterium lineatum Ehrenb., Cl. Ralfsii
Breb. var. hybridum Rabenh., Cylindroci/stis diplospora Lund, and
Penium didyrnocarpum Lund, normally produce double zygospores,
and have reason to believe in the last-named species that con-
I

jugation occurs between four cells produced by two consecutive

vegetative divisions. After carefully considering the conjugating
examples of this species which I have been fortunate enough to
obtain from several localities, I am forced to the conclusion that
there are two zygospores in close approximation, each one having
been formed by the union of a distinct pair of gametes (vide
fig. 52
D and E). In the conjugation of the two Glosteria mentioned
above, each half-cell produces a gamete, and here again there are
two zygospores each produced by the union of a pair of gametes,
one from a semicell of each plant.
The zygospore rests for a considerable time before germination.
The actual process of germination Avas first described by De Bary 1
and has rarely been observed or followed out. The two outer
coats burst and the protoplasmic contents issue in a thin vesicle

composed of an extension of the innermost coat. The contents of

this vesicledivide into two, four, or eight cells, each of which
becomes invested with a new cell-wall and gradually assumes the
form of the adult. If the species is one with characteristic

markings the first-formed cells are devoid of them, but on the first

vegetative division the new semicells acquire 'the distinguishing

ornamentation of the species. The newly formed individuals
divide repeatedly, the first few generations showing a slight in-
crease in size.

Only one true case of hybridization has been observed amongst

the Desmids and in that case the development of the zygospore
was not followed out 2 All other recorded cases of hybrids are
.

conjectural and most of them are obviously forms produced by

ordinary vegetative division.
There are many strong reasons for regarding the Desmidiaceae
as a degenerate family of Conjugata? which has originated by retro-

gression from filamentous ancestors. The degeneration has brought

1
De Bary, 'Untersuchungen iiber die Fam. der Couj.,' Leipzig, 1858.
-
Archer in Quart. Journ. Micr. Sci. 1875, pp. 414, 415. This was a zygospore
produced by the conjugation of two cells, one of which was Euastrum Didclta (Turp.)
Ralfs and the other Euastrum humerosuin Ealfs.
 Desmidiacece 143

about a loss of the filamentous condition,

accompanied by the develop-
ment of specialized morphological characters and this has gone on
1
,

hand in hand with the loss of sexual differentiation of the conjugating

cells. It has been previously mentioned that Desmidium
cylin-
dricum is the only known Desmid in which the zygospore is formed
in one of the conjugating cells
(presumably the female), and the
occasional reversion to this type of conjugation in Hyalotheca
dissiliens-goes far to prove that in all probability this was the
ancestral type of conjugation in the Desmidiacese. Moreover, it is
a type of conjugation which is represented at the
present by
Zygnemea?, although amongst the Desmids its
lingering remains
are only found in Desmidium cylindricum. A few years ago I
advocated a scheme of evolution of Desmids from ancestral fila-
mentous forms by descent through the genus Cylindrocystis 3 and ,

the recent discovery of that extraordinary member of the

Zygnemese,
Debarya Desmidioides W. & G. S. West is of surpassing interest.
4
,

This Conjugate fills up the link that was missing in the chain of
evidence which goes to show that Cylindrocystis and Mesotcenium,
and following on that nearly all the other genera of Desmids, were
most probably derived from filamentous ancestors. There is also a
great tendency towards the secondary assumption of the filamentous
condition. Not only has this resulted in the production of true fila-
mentous genera such as Spondylosiiim, Onychonema, Sphwrozosma,
Desmidium, etc., which had their origin from unicellular forms, but
this tendency reveals itself in certain
species of genera which are
normally unicellular. Thus, filamentous forms are occasionally
met with of such species as Cosmarium obliqumn Nordst. 5 C. ,

moniliforme (Turp.) Ralfs, C. Regnellii Wille Euastrmn binale

6
,

7
(Turp.) Ehrenb. and Staurastrum inconspicuum Nordst. not to
8
,

mention certain of the tropical Pleurotcenia and Micrasterias

foliacea Bail., the latter being a true filamentous form of a typically
unicellular genus with complex cell-outlines.
The Desmidiaceas is at the present day the family of Conjugates

1
W. & G. S. West in Ann. Bot. xii, 1898, pp. 53, 55.
2
Boldt in Bih. till Sv. Vet.-Akad. Handl. Bd xiii, no. 5, t. ii, f. 33 Joshua in
;

Jouru. Bot. xx, 1892.

3
G. S. West in Journ. Linn. Soc. Bot. xxxiv, 1899, pp. 409 415.
4
W. & G. S. West in Journ. Bot. 1903, p. 39, t. 446, f. 19.
5
Nordstedt in Acta Univers. Lund, ix, 1873, p. 23, t. i, f. 8.
6
W. & G. S. West in Trans. Linn. Soc. Bot. ser. 2, v, t. xv, f. 20 a'.
7
W. & G. S. West in Ann. Bot. xii, 1898, p. 30, t. iv, f. 38.
8
Borgeseu in Bot. of Faeroes, Part I, Copenhagen, 1901, p. 235, t. viii, f. 4.
144

which has attained a maximum state of specialization in the

directionof an increase in the complexity of morphological
characters, accompanied by degeneration of sexual differences.

ANCESTRAL FILAMENTOUS CONJUGATES

Genicularia \

I
Gonatozygon
t

V GONATOZYG.E
\
N

SACCODERM^E

Tetmemorus\ I

/ ^-"Ichthyocercus"Lx^
*f^
''"'
if r +S? Closterium /
/ Triploceras
j-,
Euaatrui
[Penium
^
romutum]
\. r,\.Q<Q,1VW&fc
f /'
Docidium
Pleurotaenium
\v ^___-
\

J
;
'
. Cosmocladium ;

1
Oocardiuro
I

Onyclionema
> .' 'Spondylosium
/
V ;' / Hyalotheca
\ '.
^ Sph;erozosma
'. Phymatodocis ii:lll. __. , /

\ .*** Streptonema .

.
4;
Desmidium /

^^ /
.

(Didymopn,
^ ~^ __ /
/ pLACODEBBLB
Gymnozyga x

Phytogeny of the Genera of Desmids.

There question that this complexity of outline, which is so

is little

frequently accompanied by a defensive armour of spines and spinous

processes, has been acquired as a means of defence against the
 Destt idiacece
/ 145

attacks of small aquatic animals. After the loss of the filamentous

condition it became necessary for the solitary and unprotected
individuals to acquire some other means of defence, and presumably
the present morphological complexity is the result. It is a notable
fact that those species which occur on wet rocks and in other
localities in which Amoeba?, Oligochastes, Tardigrades, Crustacea,
etc.,are either absent or very scanty, especially at high elevations,

usually possess a comparatively simple outline and are provided

with a more or less abundant mucus whereas those species oc-
;

curring in deep bog-pools, in the plankton, and the quiet margins

of deep lakes, in which localities such enemies abound, are generally

possessed of a more complicated, and in many cases of a formidable,

exterior. These characters acquired by the unicell are not only
protective against the depredations of aquatic animals, but are also
useful as anchors in the time of floods, and in their acquirement
the Law of Symmetry has exercised its full influence, with the
result that exquisite patterns have been produced which exhibit a
symmetry far ahead of that shown by any other living vegetable
organisms.
There are several thousand known species of Desmids, about
one quarter of which are British, and almost all of them can be
readily identified by their external morphological features. Some
are cylindrical with rounded or attenuated apices many of them ;

are flattened and often disciform ;

and others are of a radiating
character. To one who is not sufficiently versed in solid geometry
there are few greater surprises than the extraordinary aspects

presented by some of these plants. The majority of Desmids have

three principal axes of at right angles to one another,
symmetry
and for this reason they
require examining in three positions.
The most important aspect is the front view, in which the plant is
observed in that plane containing the two longest axes. The other
important aspects are the vertical view and the side (or lateral}
view (vide
fig.
51 E and F).
Desmids are subject to considerable variation, but only within
certain limits, and one of the most extraordinary facts relating to
these plants is the constancy of the markings embellishing the
exterior of the cell-wall. The following is a summary of the present

knowledge concerning the variation of Desmids

1
state of our :

1
G. S. West, '
On Variation in the Desmidieas and its Bearing on their Classifi-
cation,' Journ. Linn. Soc. Bot. xxxiv, 1899, p. 376.

W. A. 10
146 Chlorophycece

(1) The structure of the cell-contents is one of the most

constant features exhibited by a species but this fact can be of ;

little classificatory value owing to the very large number of

species
which possess the same structure and arrangement of the chro-
matophores.
(2) The outward form of the cell, as seen in front view, varies
within certain limits, which are usually very small, but which may
in exceptional cases (such as in pure cultures) be considerable.
The form of the vertical view is, as a rule, a more constant feature
than the form of the front view.
(3) The ornamentation (scrobiculations, granulations, spinu-
lations, etc.) of the cell- wall is relatively constant, being always

arranged according to a definite law, which is only transgressed by

variations in one or more of the individual component groups which
constitute the pattern of arrangement.
(4) The prolific growth and rapid division of immense numbers
of Desmids have a tendency to produce variations from the
typical forms.
(5) Changes in the conditions of environment cannot affect
the characters of a species unless they act for long periods of time.
It is most unfortunate that so little is known concerning the

geographical distribution of Desmids, as such a knowledge would

probably be of much greater value than one would at first sight
imagine. I have previously shown that the production of
per-
manent variation in species of Desmids under natural conditions
is much more difficult than is generally supposed 1 also, that some ;

of the prettily marked species are found in such widely separated

localities as Ceylon, Java, Hong Kong and Queensland, identical
in every respect and possessing precisely the same markings 2 It .

is likewise most improbable that this result has been

brought
about by a parallelism of modification in the course of the evolution
of these forms, owing to the occurrence of other species with a
world-wide distribution and equally constant characters. The
distribution of Micrasterias foliacea Bail, is sufficient to settle this

point, as the plant in question is the most aberrant of all forms of

v
the genus Micrasterias, having secondarily assumed a filamentous
condition and yet the plants found in North and South America,
;

India, Burmah, Siam, China, Australia and New Zealand cannot

be distinguished from one another. It should also be borne in
1
G. S. West, 1. c. p. 371.
-
W. & G. S. West in Trans. Linn. Soc. Bot. ser. 2, vol. vi, 1902, p. 124.
 Desmidiacece 147

mind that the transference by natural means of living specimens

of any Desmid from some of these countries to any of the others
is an utter impossibility, desiccation, or in many cases even partial

drying, being quickly followed by death, and submergence in

Moreover, zygospores, which might

1
sea-water equally fatal
is .

possibly withstand the entailed vicissitudes if circumstances arose

by which they could be transferred from one country to the other
(such as by the long flight of a wacling-bird), are so rarely found
that distribution by their means across an expanse of ocean is
almost impossible. There is but one conclusion to be arrived at
from a consideration of these facts, namely, that such a species has
been perpetuated by isolated communities which were derived
originally from a common stock, and that the individuals of these
communities have retained their original characters in an extra-
ordinarily constant manner. Thus, it seems probable that a sound
knowledge of the distribution of Desmids would shed much light
on the subject of previous land-connections, and might perhaps
afford more reliable evidence on this point than that afforded by
the known distribution of any other plants or animals.
As a significant instance, it may be mentioned that several

remarkably Desmids, such as Staurastrum Ophiura Lund., St.

fine
Arctiscon (Ehrenb.) Lund., St. jaculiferum West, St. longispinum
(Bail.) Arch., St. Cerastes Lund., St. Brasiliense Nordst. var. Lundellii
W. & G. S. West, Micrasterias furcata Ag., M. conferta Lund.,
Pleurotcanium nodosum (Bail.) Lund., and others, are not uncommon
in the eastern parts of N. America and that in Europe they are

only abundant on the north-western shore districts of Ireland,

Wales, Scotland, the Outer Hebrides, Scandinavia and Lapland.
One of these species Staurastrum jaculiferum West is also
known from the Shetland Is., the Faeroe Is. and Iceland.
Borge has examined a number of subfossil Desmids from the
glacial clays of the Isle of Gotland, and a few subfossil forms of
existing species have also been noted from an ancient peat deposit
near Filey in E. Yorkshire.
Desmids thrive best in soft water, and they are most numerous
in peaty water which has a trace of acidity. With few exceptions
they do not flourish in water containing carbonate of lime in
solution, and no good collection of Desmids can be made in a
purely limestone district in which the water is hard.

1
One Desmid has been described as inhabiting brackish water, viz.: Cosmarium
saliniun Hansg. in Oesterr. bot. Zeitschr. 1886, p. 335.
102
148 CJdorophycece
I have attempted to give a reasonable classification of the

genera of Desmids based upon the evolutionary scheme previously

mentioned 1
and upon the recent and sound observations of
Liitkemtiller 2 It is very intere3ting to note that Liitkemiiller's
.

conclusions, which he arrived at chiefly from the study of the

minute structure of the cell-wall, coincide almost entirely with the
scheme I put forward in 1899, which was based upon a compre-
hensive study of the external morphological features of these plants.
The division of Desmids into filamentous forms and solitary
forms, such as is adopted by many writers even at the present

time, no longer tenable and such an arrangement as that given

is ;

3
by Bessey in which he places Desmids under the three tribes
,

'

Desmidiese,' Arthrodieas and Cosmarieae,' is obviously one which

' ' '

is not based upon a careful study of the Desmids themselves

and is therefore of no value.

Most Desmids would appear to have had an origin from the
genera Mesotcenium and Cylindrocystis, which were themselves
4
derived by retrogression from ancestral filamentous Conjugates .

The genera Gonatozygon and Genicularia have so little in common

with most other Desmids, and resemble so much some of the
present existing filamentous Conjugates, that they must be re-
garded as having had a distinct origin from filamentous ancestors.
They are however, more specialized than the genera Spirotcenia,
Mesotcenium or Cylindrocystis.
The genus Penium is the most difficult to relegate to its proper
place. In the sense in which this genus is generally regarded ib
undoubtedly contains many widely different plants, and therefore
one hails with delight Liitkemuller's suggestion that the name
Netrium be again brought forward as a distinct genus to include
four species which obviously form a natural group. On the removal
of these four plants from the genus, the remainder do not fit
satisfactorily into any one place in the scheme of classification,
and the difficulty lies in the fact that our knowledge of the plants

1
G. S. West in Journ. Linn. Soc. Bot. xxxiv, 1899.
2
Liitkemtiller in Beitriige zur Biol. der Ptianzen, herausgegeben von Dr F. Cohn,
Bd viii, Breslau, 1902.
Bessey in Trans. Amer. Micr. Soc. xxii, I'.lOl.
3
4
Had these genera originated directly from Flagellate forms, it is scarcely
conceivable that the motile condition would not be frequently reverted to in fact,
;

it is reasonable to suppose that it would play a considerable part in the life-history

of these Algae. Such is the cose in /ill ijronps of Alr/tc with it direct Flagellate
ancestry and the entire absence of motile forms of any description from the
;

Desniidiaceas lends strong support to the view put forward from other considera-
tions, of the origin of this family of unicells by retrogression.
 Desn iidiacece 149

istoo scanty to allow of splitting them up into a number of separate

genera. One cannot see how, in the present state of our know-
ledge, such genera could be denned, and yet some of these plants
are certainly not nearly related. The plant commonly known as
Penium minutum (Ralfs) Cleve is the most puzzling of all.
I have accepted the two sub-families proposed by Liitkemuller
"
and also his five tribes, but have placed the " Gonatozygae first
" "
and altered the definition of the Penieaa in order to include the
heterogynous collection of Desmids embraced in the present genus
Penium.
The following is a synopsis of all the known genera of Desmids,
four of which (Ichtkyocercus, Triploceras, Phymatodocis and Strep-

tonema) are exclusively tropical and one (Ancylonema) is exclusively

arctic.

Sub-family I. SACCODERM.B.
Cell-wall uusegmented and without pores. Point of division of cells
indefinite and unknown previous to the actual division. The young half of
the cell is developed obliquely and its walls are absolutely continuous with .

the walls of the older half.

Tribe 1. G-onatozygae. Cells elongate, cylindrical and unconstricted,

forming loose filaments. Cell-wall with a differentiated outer layer of which

the small roughnesses and spines form a part.
*
Chloroplasts axile 1. Gonatozygon.
** and twisted ... 2. Genicularia.
Chloroplasts- parietal spirally
Tribe 2. Spirotaeniese. Cells solitary, relatively short and mostly un-
constricted. Cell-wall a simple sac, without a differentiated outer layer.
The cell becomes adult by periodical growth.
* One
chloroplast in each cell.
t Chloroplast .spirally twisted, axile or parietal 3. Spirotcenia.
tt Chloroplast plane, axile.
Cells solitary 4. Mesotcenium.
Cells forming short filaments f>.
Ancylonema.
** Two chloroplasts in each cell.

t Chloroplasts star-shaped, radiating from a

central pyrenoid 6. Cylindrocystis.
tt Chloroplasts ridged with longitudinal
serrated ridges
V-v . 7. Netriwm.

Sub-family II. PLACODERM.E.

Cell-wall segmented, with a differentiated outer layer. Cell-division follows
a fixed type, with the interpolation of the younger halves between the old
ones. The younger portions of the cell-wall are joined to the older portions
by an oblique surface.
150 Chlorophycece
A. Point of division of cells variable or sometimes fixed (at the isthmus).

Tribe 3. Peniese. Cells of moderate length, straight, cylindrical, some-

times with a slight median constriction. Cell-wall with or without pores.
Point of division of cells often variable. The cell often arrives at maturity
by periodical growth.
8. Penium.

Tribe 4. Closteriese. Cells elongate, generally curved symmetrical in ;

one longitudinal plane only. Cell-wall commonly with pores. Points of

division regularly placed in the middle region of the cell.
* Cells almost cylindrical, scarcely attenuated.
Chloroplast single, without apical moving
9. Roya.
grannies
** Cells
strongly attenuated towards each ex-
tremity. Two chloroplasts in each cell,
with apical moving granules 10. Closterium.

B. Point of division of cells always fixed (at the isthmus).

Tribe 5. Cells exhibit great variety of form, and the cell-

Cosmarieae.
wall consists of two thin, firm layers with pores. There is no periodical
growth, the cell becoming adult immediately after division by the mature
growth of the young semicell.
a. The point of division of the cell, where the new and old parts of the
cell-wall are obliquely fitted together, remains plane during division. Solitary
or colonial.
* After division the cells remain free and solitary.
t Cells elongated and cylindrical constriction slight.
;

Apices of cells truncate or rounded.

\ Base of semicells plicate 11. Docidium.
H Base of semicells plane 12. Pleurotcenium.
Apices of cells cleft, incision open or narrow.
| Cell-wall adorned with rings of
furcate processes 13. Triploceras.

II Cell- wall plane.

Apical incision widely open,
each apical angle furnished
with a spine 14. Ichthyocercus.

Apical incision narrow ... 15. Tetmemorus.

tt Cells relatively short, commonly compressed or radiating, con-
striction usually deep.
Cells compressed (at right angles to the plane of the fr< >nt
view) from the vertical view fusiform or elliptical.
;

I Cells generally with an apical in-

cision and a central protuber-

ance, moderately lobed 16. Eua strum.

II Cells very compressed and deeply

lobed or incised 17. Micro sterias.

Cells with a more or less entire margin, often fur-

nished with warts or spines.
 Desmidiacece 151

Cells commonly with a central protuberance.

IF Cell-wall smooth, granu-
late, verrucose, etc.
Central protuberance
present or absent 18. Cosmarium. /

TTTT Cell-wall with regularly

arranged spines, com-

monly in pairs. Cen-
tral protuberance al-

ways present 19. Xanthidium.

Cells without a central pro-
tuberance; angles spinate 20. Arthrodesmus.
Cells from the vertical view com-

monly radiating, triangular, quad-

rangular, or up to 11 -radiate, rarely
fusiform 21. Sta vmstrum. *
** After division the remain attached to form colonies.
cells

t Colonies spheroidal cells not in contact, but joined by gelatinous

;

bands.
Gelatinous bands narrow; few cells

forming a microscopic colony 22. Cosmocladium.

Gelatinous bands very broad; many
cells forming a macroscopic colony 23. Oocardium.
ft Colonies thread-like; cells attached by their apices to form
long filaments.
Cells attached by special apical processes. _,

i Apical processes very short 24. SpJuerozos/itn.

1| Apical processes long and over-

lapping the apices of the ad-
joining cells 25. OnychoneiiKi.
Apices of cells plane and flat.

\ Cells deeply constricted.

Cells in vertical view elliptical 26. Spondylosium.
Cells in vertical view quad-

rangular with produced

angles 27. Phymatodocis.
\\ Cells very slightly constricted... 28. Hyalotheca. \/
b. The point of division of the cell, where the new and old parts of the
cell-wall are obliquely fitted together, develops a girdle-like thickening or

ingrowth, which projects both ways into each of the old semicells during
division. Cells attached to form thread-like colonies.
* Cells 29. Streptonema.
joined by special apical processes
** Cells
joined by their flat apices or by flattened apical projections.
t Cells short, in vertical view fusiform, trian-

gular or quadrangular (rarely circular

with produced angles) 30. Desm.idium.-y

tt Cells elongate, cylindrical 31. Gymnozyga.

152 CKLorophycece

Sub-family I. SACCODERM^E.
This a very natural group of Desmids in which the cell-wall
is

is unsegmented and destitute of pores. There is no line of de-

marcation between the
newer semicell and the
older semicell, the cell-
wall being absolutely
continuous. Division
takes place at no fixed
part of the cell (except
in a few species of

Cylindrocystis)a,nd the
young semicells are
developed obliquely,
gradually sliding away
from one another as
they reach maturity.

Tribe 1. Gonatozygce.
This tribe only
includes two genera
which are considerably
removed from other
Desmids. The cells
are long and cylindri-
cal, and are joined by
their apices into fragile
filaments of variable

length. The filaments

easily break up, but
Fig. 53. A and B, Gonatozygon Brebissonii De
this in no way inter-
Bary x 464) A, from Esher Common, Surrey;
( ;

B, from Streusall, N. Yorks. C E, G. Brcbixxonii feres with the life of

var. lave (Hilse) W. & G. S. West, from Mickle
F and G, G. Bn'bixxoiiii the individual cells.
Fell, N. Yorks. ( x 356).
var. minntum W. & G. S. West, from Kiccall
Conjugation only takes
Common, E. Yorks. ( x 464). H, zygospore of G. between cells
mouoteenium De Bary ( x 464). I and J, Gcnicularia place
Rpirotania De Bary, from near the Lizard, Cornwall; which have become
I, vegetative cell ( x 312)
; J, zygospore ( x 220).
free. The cell-wall is

differentiated into two layers, the inner layer being hyaline and
 Desmidiacece 153

structureless, the minute prominences or small spines characteristic

of most of the tribe forming a part of the outer layer.
Genus Gonatozygon De Bary, 1856. The cells are cylindrical
or narrowly fusiform, 10 20 (rarely 40) times longer than the
diameter, and without any constriction. The apices are occasion-
ally subcapitate and always more or less truncate. The apical
attachment of the very slight, the least disturbance causing
cells is
a general dissociation of the filament. The cell-wall is rarely
smooth, being usually covered with minute, sharp prominences or
short spines. There are generally two axile chloroplasts in each
cell (rarelyonly one), each of which is rather narrow, undulated,
sometimes twisted, and contains 4 16 equidistant pyrenoids.
Occasionally an apical vacuole containing moving granules is
present at each end of the cell, beyond the limit of the chloro-
plasts. On the formation of the zygospore the conjugating cells
frequently become geniculate. The zygospore has a smooth outer
coat.
There are five British species of De Bary
the genus, of which G. monotcenium
[G. Ralfsii De Bary] and G. Brebissonii De Bary
Docidium asperum Breb.]
[?

(fig. 53 A
and B) are the most abundant. Smooth varieties of both these
species are known and G. Kinahani (Arch.) Rabenh., which is the largest
species of the genus (length of cell 162 376 p. breadth 11 14 /*),
;
is always
smooth. The smallest form is G. Brebissonii var. minutum W. & CI. S.

West (fig. 53 F and G), which has a length of 47 '5 67 5 ^ and a breadth of
-

4-27 p.

Genus Genicularia De Bary, 1858. The cells are similar in

form to those of Gonatozygon, being cylindrical, elongate, uncon-
stricted and with truncate apices. The filaments are extremely
fragile and the cells always become free previous to conjugation.
The zygospore is globose and smooth, and the conjugating cells
become geniculate. The cell-wall is rough on the exterior almost
exactly like the cells of Gonatozygon monotcenium. There may be
two or three parietal chloroplasts in each cell, disposed as spiral
bands or rarely somewhat irregular. Each chloroplast contains a
number of pyrenoids. Genicularia is one of the rarest known genera
of Desmids.
G. Spirotcenia De Bary has only been found in the British Islands from
Cornwall and the Shetland Is. length of cells 200400 ^, breadth 2025 p.,
;

diam. zygosp. 48 57 p (fig. 53 I and J). G. elegans W. & G. S. West is a

more slender species in which the chloroplasts form laxer spirals length ;

303 427 p., breadth 14 16'3 p.. It is only known from the plankton of Loch
Fadaghoda, Lewis, and of Loch nan Fun, N. Uist, Outer Hebrides.
154 Chlofophycece

Tribe 2.
Spirotceniece.
In the British genera of this tribe the cells are solitary,
relatively short and unconstricted (with the exception of a few
species of Cylindrocystis}. The cell- wall has no differentiated
outer layer and is quite smooth. The individuals reach the adult
condition by periodical growth, chiefly in length.
Genus Spirotaenia Breb., 1848. The cells are straight or very
slightly curved, oblong-cylindrical or fusiform, and frequently
enveloped in mucus. There is no median constriction and the
apices of the cells may be rounded, truncate, subacute or very
acute. There is only one chloroplast in each cell, which may be
band-like and parietal, or axile and cristate, and is always spirally
twisted to the left. The nucleus is generally asymmetrical, and
the cell-wall is smooth and structureless. The genus is divided
into two sections; in sect. 1, Monotceniece Rabenh., the chloroplast
is a parietal band spirally arranged round the inside of the cell-
wall in sect. 2, Polytceniew Rabenh., the chloroplast is axile with
;

a variable number of spirally twisted ridges. There may be one

or many pyrenoids in the chloroplast.

There are fourteen British species of the genus, none of which is common.
S. condensata Breb. (fig.
54 A ; length 150270 p.,
breadth 1827 /x)
is the

largest and most widely distributed, and /S'. closteridia (Breb.) Arch, is the
smallest (length 13'5 p., breadth 4'5 4'6 p.). They are all very delicate, with
thin cell-walls, and most of them can only be identified with certainty from
living specimens. The zygospores of few of them are known and they are
rarely met with the outer layer of the
;
wall of the zygospore is usually
ornamented. Most of the species occur in peat-bogs.

Genus Mesotsenium Nag., 1849. The cells are cylindrical or

subcylindrical, generally straight or slightly curved, and are

with-
out any trace of a median constriction. The apices are, as a rule,

broadly rounded. The chloroplast, of which there is usually only

one in a cell, is a flattened, axile plate extending from end to end
of the cell, and there may be one or several pyrenoids. Sometimes
there are two flattened chloroplasts. The cells often contain
numerous globules of an oily nature.

There are ten British species of the genus, seven of which occur as
mucilaginous masses amongst mosses and hepatics, generally on wet rocks.
The largest of these is M. De Greyi Turn. (fig. 54 U; length 74 125 p.;
breadth 15 '5 30 /a) and the smallest is M. macrococcum (Kiitz.) Eoy & Biss.
var. micrococcum (Kiitz.) W. & G. S. West (length 13'5 15'3 p. ;
breadth
 Desntidiacece 155

8'6 /i). M. caldariorum (Lagerh.) Hansg. is an attenuated species which

occurs in greenhouses, forming a thin mucilaginous stratum on damp walls,
etc. (length 27 46 fi breadth 1O5 11-5/t). M. Endlicherianum Nag. is the
;

most frequent of the free-floating forms (length 25 27 breadth 8 -5 9 5 /*),

/LI ;
-

and M. purpureum W. & G. S. West and M. molascens De Baiy contain

phycoporphyrin. The zygospores of species of this genus are generally poly-
hedral with thick brown walls.

n nx<

Fig. 54. A, Spirottenia condensata Breb., from Rhiconich, Sutherland ( x 334),

showing parietal chloroplast. B, 8p. obscura Ealfs, from Terrington, N. Yorks.
(
x 435), showing the axile, cristate chloroplast. C, zygospore of .S'/>. trnncata
Arch. (
x 250, after Archer). D, Mesotcenium De Grci/i Turner, from near
Settle, W. Yorks. ( x 435). E and F, M. macrococcum (Kiitz.) Roy & Biss.,
from near Giggleswick, W. Yorks. ( x 334). G, zygospore of M. chlamydosporum
De Bary, from Mayo, Ireland ( x 334). H and I, Ct/lindroci/atix Brebissonii
Menegh., from Helvellyn, Westmoreland; H, vegetative cell; I, zygospore
(
x 435). J, Cyl. diplospora Lund., from Galway, Ireland ( x 435). K, Netrium
Digittts (Ehrenb.) Itzigsh. & Rothe, from Moel Siabod, N. Wales ( x 435).

Genus Cylindrocystis Menegh., 1838. The cells are -straight

and cylindrical, averaging twice longer than the diameter, and
they niay or may not possess a slight median constriction. The
156 Chlorophycece

apices of the cells are generally rounded or truncately rounded.

There is a single axile, substellate chloroplast in each semicell and
in the centre of each chloroplast is a large pyrenoid. The radiating

prolongations of the chloroplast vary in number in the different

species, and often become flattened against the interior of the
cell-wall.

The most abundant species is Cyl. Brebissonii Menegh. (length 43 55 p.;

breadth 15 18 /x, fig. 54 H and I) which occurs in quantity in upland

Sphagnum-bogs and in peaty pools. It sometimes occurs in pure masses both

amongst Sphagnum and on wet peat. Cyl. crassa De Bary also occurs in
Sphagnum-bogs and amongst other Alga and mosses on wet rocks. Cyl.
1

diplospora Lund, prefers the waters of lakes and quiet pools, and is a much
rarer plant one form of it var. major West is the largest member of the
;

genus; length 102 114 p.; breadth 48 54 p..

Genus Netrium. (Nag., 1849). genus with straight, cylin- A

drical, oblong-cylindrical or fusiform cells, and without any median
constriction. The chloroplasts are two (in one species four) in
number, one (in one species two) in each semicell each chloro- ;

plast axile with about six radiating longitudinal plates.

is These
plates are deeply notched along their free edges in all except
Netrium interruption. There are several pyrenoids in each chloro-
plast, generally smalland scattered. The plants placed in this
genus are excluded from the Placoderm Desmids, and therefore
from the genus Penium, on account of the structure of their cell-
wall, which is
unsegmented, without pores, destitute of a differen-
tiated outer layer, and quite smooth.
N. Digitus (Ehrenb.) Itzigsh. & Rothe is one of the most abundant
Desmids of elevated boggy moorlands and varies much in form and size ;

length 130 387 p. breadth 40 82 /x fig. 54 K. N. oblongum (De Bary)

; ;

Liitkem. occurs in similar situations but is not quite so frequent length ;

96 135 p. breadth 32 33 p.. N. interruptum (Breb.) Liitkem. is cylindrical

;

with obtusely conical apices, and each chloroplast is transversely segmented,

so that there are four chloroplasts in each cell arranged in an axile series. In
this species the free edges of the plates are not notched length 220 320 p. ; ;

breadth 3764 M .

Sub-family II. PLACODERM.E.

This group includes the great majority of Desmids and is
characterized by the segmented cell-wall with its differentiated
outer layer. The cell-wall usually possesses pores, but this is not
invariably the case. There is
always a very distinct line of demarca-
tion between the old and new semicells, the younger portions of
 Desmidiacece 157

the cell-wall not being continuous with the older portions, but
joined to them by a narrow, oblique or bevelled surface. The cell-
division is of a fixed type, taking place strictly in the manner of

previous cell-divisions, and the younger semicells are interpolated

between the older ones. The sub-family can be divided into two
which includes those Desmids in which cell-division
sections, one of
does not always take place at a fixed point and those in which it
does always take place at a fixed point known as the isthmus.

A. Point of division of cells variable or sometimes fixed

at the isthmus.

Tribe 3. Peniece.

This tribe only includes one genus, in which the cells are

solitary,of moderate length, straight and generally cylindrical.

Sometimes, but not always, there is a slight median constriction.
The points of division are often variable, although the actual cell-
division is of thesame type. The cell-wall may be with or without
pores, and the cell often grows periodically until it becomes adult.
Genus Penium Breb., 1844. This genus is difficult to strictly
define, as it undoubtedly contains many species which will ulti-

mately have to find a resting-place elsewhere. The cells are

straight, cylindrical, subcylindrical, ellipsoidal, or fusiform, and the

apices may be rounded or truncate. The vertical view is always
circular.The inner layer of the cell-wall is in many forms orna-
mented with striations, punctulations or granulations, but in others
the cell-wall is quite smooth. There is one axile chloroplast in
each semicell, with radiating longitudinal plates which have the
free margin entire. The pyrenoids are one or many and uniseriate.
Sometimes there is a prominent vacuole near each extremity of
the cell containing moving granules.
There are about 27 British species of Penium, of which P. Libellula (Focke)
Nordst. (length 250354 p, breadth 3851 p. fig. 55 D) is one of the largest
; ;

and most striking. P. margaritaceum (Ehrenb.) Brt-b., P. CyUndrus (Ehrenb.)

Breb. (fig. 55 A and B) and others, are granulated species, and P. spirostriolatu/n
Barker (length 123274 p. breadth 23 26 p. fig. 55 C) is a species with spiral
; ;

striations on the cell-wall, which often anastomose or become broken up into

dot-like thickenings. P. polymorpltum Perty (length 55 58 p.; breadth 25
27 p.) is one of the most general of the upland, moorland species, and some of
the smallest species of the genus are P. iiiconspieuum West (length 14'5 19 p.;
breadth 5 5'7 p.), P. minutissimum Nordst. (length 12 2 -
13 p.; breadth
158 Chlorophycece
6-87-3 p.)
and P. suboctangulare West (length 10'7 11'5/*; breadth 6'8
7 /* ; fig. 55 E). The cell-wall of many of the species is of a yellow or reddish-
brown colour. P. minutum
(Ealfs) Cleve is relatively the
longest species of the genus,
but should, perhaps, be rele-
gated elsewhere.

Tribe 4. Closteriece.

The cells are elongate,

generally curved, and usu-
ally attenuated towards
each extremity. There is
no trace of a median con-
striction and the plants are
circular in transverse sec-
tion. The points of division
are always in the middle

region of the cell and the

cell-wall usually possesses

pores. The cells are only

symmetrical in one longi-
tudinal plane.
Genus Roya West &
G. S.
West, 1896. This
genus was named after the
late Dr John Roy, who first
pointed out the differences
between those plants which
are now included in it and
55. A and B, Penium Ci/limlm*
species of Closterium. The
Fig.
(Ehrenb.) Breb. ; A, from Loch Minnoch,
Kirkcudbright; B, zygospore from Thursley cells are elongate, cylindri- .

Common, Surrey ( x 474). C, P. spirostrio-

latum Barker, from Kerry, Ireland (x474). cal, very slightly curved and
D, P. Lilelliila (Focke) Nordst., from Bowness, scarcely atten uated towards
Westmoreland ( x 200). E, P. suboctangulare
the poles. There is
only
West, with zygospore, from Killarney, Kerry,
Ireland ( x 365). F, P. curt-urn Breb., from one chloroplast which occu-
Grimspound, Devonshire ( x 474). G and
H, Kui/u obtum (Breb.) W. & G. S. West pies almost the entire cell-
var. montana W. & G. S. West, from Baildon
cavity. It extends to within
Moor, W. Yorks. ( x 570). I, R. C<i>ni-ic<i
W. & G. S. West, from Llyn Ogwen, N. Wales a very short distance of
(x474). J and K, E. Pseudoclosterium (Roy) each pole and its extremi-
W. & G. S. West, from Pilmoor, N. Yorks.
ties are convex. In the
(x474).
 Desn lidiacece 159

median portion of the cell there is a slight lateral excavation in

the chloroplast for the lodgement of the nucleus, which is thus
asymmetrical. There are from four to thirteen pyrenoids in the
chloroplast, arranged in a single series. The cell-wall is
relatively
thin and without pores.
The genus is readily distinguished from Glosterium by the
absence of any marked attenuation towards the apices and by the
single chloroplast with convex extremities, the latter being in
such close proximity to the ends of the cell that there is no room
for an apical vacuole. The lateral position of the nucleus is also
characteristic.

There are only three species of the genus, all of which occur in Britain.
R. obtusa (Breb.) W. & G. S. West (and its var. montana; fig. 55 G and H)
is not an uncommon Desmid in mountainous districts; length 48 117 /x;
breadth 5'5 12'5 p.. R. Pseudoclosterium (Roy) W. & G. S. West is a very
narrow species of rare occurrence; length 98 192 p.; breadth 2 6 3 p. -

fig. 55 J and K. R. Cambrica W. & G. S. West is only known from N. Wales;

length 173 177 p.;
breadth (5'2
-
6 7 p.; fig. 55 I.

Genus Clostermm Nitzsch, 1817 The cells are elongate,

l
.

generally distinctly curved, and often markedly lunate or arcuate.

In most cases they are strongly attenuated towards the poles, the
latter being obtuse, truncate, rostrate, or drawn out into long
needle-like points. In most of these plants the cell-wall possesses
pores and in a large number of species it is striated, the striations
being internal thickenings of the cell-wall running from pole to
pole and disposed at regular intervals. The number and strength
of the striations varies very much in different species of the

genus. There are two chloroplasts, one in each semicell, and

there may be one or many pyrenoids in each chloroplast. The
extremities of the chloroplasts are concave and they do not reach
the apices of the cell, thus affording room for prominent apical
vacuoles which contain one or many moving granules of variable
size In those species in which the apices are greatly
and shape.
produced the chloroplasts dd not extend into the prolongations,
1
The name "Arthrodia" (Raf. in Desv. Journ. 1813, i, p. 235) cannot possibly
be utilized as a generic name in place of Clo&terium. Rafiuesque's diagnosis
applies equally to Aiikixtrodesmus, Penium, Docidiuin, Pleurotanium, Cylindroeystis,
and Ti'tmemorus. Nordstedt (in Hedwigia 1893, Heft 3, p. 148) has clearly pointed
" "
out that " Arthrodia" must always remain a "genus ignotum and a nomen deleu-
durn." The same author's remarks should also be consulted concerning "(;////>.-,
"
Ehreub.," "HeUerella Bory," "Ursinella Turpin," Prolifera Vauch." and"Cora-
jiujata Vauch."
160

and the apical vacuoles are often correspondingly lengthened out.

The chloroplasts are similar in form to those of the genus Penium.
consisting of an axile mass with more or less distinct longitudinal

Fig. 56. A, Closterium acerosum (Schrank) Ehrenb.,

from Glen Shee, Perthshire
(x200). B Cl. striolatuni Ehreub., from Frensham, Surrey; B and C,
D,
outlines, x 125 D, x 260. E, Cl. acutinn Breb., from Sligachau, Skye ( x 474).
;

x
F, Cl.parvulum Nag., zygospore, from Esher West-end Common, Surrey ( 474).
G, CL rostratum Ehrenb. var. brevirostratum West, zygospore, from Wimbledon
Common, Surrey ( x 200).

ridges. In Cl. acutum Breb. and several others there are no

longitudinal ridges.
The curvature of many of the species of this genus is very
constant and may be utilized as a specific character. In stating
 Desmidiacece 161

the measurements of a species, the diameter across the middle of

the cell should be given, the distance between the apices, and the
curvature of the outer margin expressed in degrees of arc.
The zygospores are generally globose and smooth, but the
spherical zygospore of Cl. co.losporum Wittr. is furnished with
conical papillations. The zygospores of certain species, such as
Cl. rostratum Ehrenb. (fig. 56 G)
and Cl. Kutzingii Breb., are
flattened and quadrate with truncate angles that of Cl. Venus ;

Klitz. is also angular and sometimes twisted.

There are about 60 British species of the genus, of which Cl. turgidum
Ehrenb. (length 476 940 p. breadth 43 80 /*) is the largest and Cl. pusillum
;

Hantzsch var. monolithum Wittr. (length 29'8 40'4 breadth 7'5 8'6 //,)
,u. ;

the smallest. Cl. aciculare Tuffen West and C. pronum Breb. are the most

elongate species of the genus, and certain forms of Cl. acutum Breb. (fig. 56 E)
are the narrowest. The commonest and most widely distributed species are
Cl. parvulum Nag. (fig. 06 F), Cl. Venus Kiitz., Cl. Leibleinii Kiitz. (fig. 51 B),
Cl. moniliferum (Bory) Ehrenb., Cl. Ehreribergii Menegh. (fig. 51 A) and Cl.

acerosum (Schrank) Ehrenb. Cl. gracile Breb. is often abundant in Sphagnum

bogs. The commonest striated species are Cl. striolatum Ehrenb. and Cl.
rostratum Ehrenb.
Liitkemiiller has divided the genus and I think quite correctly into two
groups, the one being characterized by the possession of an elongated
first

median girdle, which is an intercalation of a cylindrical piece of cell-wall

resulting from a growth to maturity subsequent to cell-division. The second
group is characterized by the absence of this girdle, the two daughter-cells
becoming adult immediately after cell-division.

B. Point of division of cells always fixed at the isthmus.

Tribe 5. Cosmariece.

The great majority of Desmids are included in this tribe and

amongst them there is great diversit/ of form and size. They are
solitary or filamentous, or grouped in spherical colonies, and most
of them possess at least three planes of symmetry. The cell-wall
consists of two thin, very firm layers, with pores, and is frequently
ornamented with spines, warts and excrescences of all kinds.
There is no periodical growth, the cell becoming adult immediately
after division by the mature growth of the younger semicell.
Series a. The oblique junction of the new and old halves of
the cell-wall (at the region of the isthmus) remains quite plane
during division. The plants are solitary or colonial.
W. A. 1 1
102 Chlorophyced'
Genus Docidium Breb., 1844 em. Lundell, 1871. The cells
;

are comparatively small,

n elongate, straight and
slightly constricted. They are subcylindrical
with an inflation on each side of the constric-
tion, or nodulose from pole to pole. The base
of each semicell is plicated and there is a
small basal granule under each plication. The
basal plication of the semicells is the
principal
point of distinction between this genus and
Pleurotceniam. The apices of the cells are
always truncate and smooth. There is a central
chloroplast in each semicell, very irregular in
form, and containing an axile row of pyrenoids.
The zygospores are unknown.
There are three British species, of which D. Bacidum
Breb. is the. only one that
widely distributed, and
is

even it is distinctly rare; length 167 262 p. breadth ;

at basal inflation of semicells 12 13 p; tig. 57 C. A

D. undulatum Bail, is restricted to the western
districts of Ireland and Scotland, and the semicells
Fig. 57. A C,Do-
cidinmBaculumSrel)., have a nodulose appearance caused by repeated shallow
from Bowness, West- constrictions from base to apex length 187
;
262 p. ;

moreland A, x 315 ;
breadth at basal inflation of semicells 13 16 p.
fig. 57
;
;

B, base of serny:ell,
x 630 C, basal view
;
D and E.
of semicell, x 630.
D and E, D. undit- Genus Pleurotsenium Nag., 1849: The
latum Bail., from and slightly con-
ce }i s are elongate, straight,
near Oughterard, Gal-
way, Ireland D, ; They are cylindrical, with or without
stricted.
x 315 E, base of
;
on either side of the constriction, or
inflations
semicell, x 630.
they may be nodulose along their entire length.
They are slightly attenuated towards each pole and the apices
are usually truncate, often being furnished with a ring of conical,
tooth-like projections. There is no basal plication of the semi-
cells. The chloroplasts are disposed as longitudinal, parietal bands,
several in each semicell, and are furnished with a number of
pyrenoids. Sometimes they become broken up into numerous
parietal pieces, each containing a single pyrenoid. In most of the
species the central portions of the cell contain large vacuoles and
occasionally numerous moving granules are observed in the terminal
or subterminal ones. These granules are of a yellow colour and
black when
seen in mass, differing in this respect from the moving
granules normally present in the genus Closterium. They are
 Desmidiacece 103

produced mostly by pathological conditions. Zygospores of few

species are known they are globose and smooth.
;

Fig. 59. Tetmemorim tjranulatus

(Breb) Kalfs, from Lough Anna,
Donegal, Ireland. A, front view of
cell ; B, zygospore (
x 315).

Fig. 58. A, Pleuroteenium coronatum

(Breb.) Eabenh., from Helvellyn, West-
moreland ( x 236). B, zygospore of PI.
Ehrenbergii (Breb.) De Bary, from
Thursley Common, Surrey ( x 315).

Species of this genus are much more abundant in the tropics than in
temperate regions, and the tropical species frequently assume a secondary
filamentous condition. Only nine species are known as British, of which
Pi. maximum (Reinsch) Lund, is the largest, reaching a length of 852 p,
and
a diameter of 54 p..
The most abundant species is PL Ehrenbergii (Breb.)
De Bary; length 240 400> breadth 18 24,*; fig.
;
58 B. PL Trabecula
(Ehrenb.) Nag. and PL truncatum (Breb.) Nag. are each widely distributed.
PL nodosum (Bail.) Lund, is the most striking species and is confined to the
western districts of Wales, Ireland and Scotland.

Genus Tetmemorus Ralfs, 1844. This is a well-marked genus

with straight cells of moderate length, slightly constricted in the

112
164 Chlorophycece

middle and deeply cleft at each apex. The cells are usually
attenuated towards each pole and the apical cleft or incision is
tightly closed. There is a central chloroplast in each semicell
with a single axile row of pyrenoids. The zygospores are globose
and smooth, or subquadrate with rounded angles.
There are only four British species, T. granulatus (Breb.) Ralfs (length
138238 p.; breadth
3045 /* fig. 59) and T. Icevis (Kiitz.)
;
Ralfs (length
57 76/u,; breadth 19 25 being the most widely distributed.
ju.)
The former,
which possesses a sparsely scrobiculated cell-wall, is one of the most ubiqui-
tous of Desmids and the latter has a distinct preference for mountainous
districts. T. Brebissonii (Menegh.) Ralfs is a more or less frequent Desmid
in bogs and pools containing submerged Sphagnum.

Genus Euastrum Ehrenb., 1832. In this genus the cells are

relatively shorter than in any of the preceding genera of the

Fig. 60. A, Euastrum elegnns (Breb.) Kiitz. (a form), from Capel Curig, N. Wales
(
x 474). B, zygospore of E. elee/ans, from New Forest, Hants. ( x 474). C, E.
Mimic (Turp.) Ehrenb., from Thursley Common, Surrey ( x 474). D, zy^o-
spore of E. oblontjinn (Grev.) Ralfs, from Pilmoor, N. Yorks. ( x 200). /, front
view s, side or lateral view c, vertical view.
; ;

Cosmarieaa and they are always distinctly flattened. Seen from

the front view they are elliptical, or narrowly elliptical, and they
possess a median constriction of considerable depth. The lateral
margins entire or lobed, and the apices are usually deeply
may be
notched. In the lobed species there is always an odd number of
lobes to each semicell, the apical lobe (which bears the notch)

being termed the 'polar lobe.' In the vertical and side views
 Desu iidiacece 165

there is generally a well-marked protuberance in the middle of

each side of the semicells. There is one axile chloroplast in each
semicell, which is often very irregular; occasionally it
possesses
irregularly disposed, longitudinal plates. There
a single central is

pyrenoid in the small species and several scattered ones in the

large species. The zygospores are globose or ellipsoid, and are
furnished either with numerous simple spines or with conical
papilla?.
There are 43 British species of the genus, about half of which are widely
distributed. The smallest and perhaps the most abundant species of the
genus is E. binale (Turp.) Ralfs; length 10 20 /z; breadth 9 16 ju.; thickness
5-5 7 /A; fig. 60 C. E. elegans Breb. (fig. 60 A and B), E. ansatum Ehrenb.,
E. Didelta (Turp.) Ralfs and E. oblongum (Grev.) Ralfs (fig. 60 D) are widely
distributed all over the country, but in some districts they are by no means
common. E. insigne Hass. (length 98 110 /j.
breadth 49 60 ^) is an upland
;

form found abundantly amongst Sphagnum. E. verrucosum Ehrenb. and

E. gemmatum Breb. are two of the prettiest species, and E. crassum (Breb.)
Kiitz. (length 167 185 ^; breadth 87 97 p) and E. oblongum (Grev.) Ralfs
(length 144 167 // breadth 80 83 /i) are the largest species found in Britain.
;

E. pectinatum Breb. is more frequently found with zygospores than any other
species of the genus.

Genus Micrasterias Ag., 1827. [Holocystis Hass., 1845 ;

Tetrachastrum Dixon, 1859.] This genus contains some of the

largest and prettiest of Desmids. The cells are flattened, in many
cases almost disc-shaped, and they are circular or broadly elliptical
in outline. The semicells are divided by deep incisions into either
three or five lobes, of which the polar lobe may be entire or
furnished with a wide excavation at the apex. The lateral lobes
are sometimes narrow and attenuated, but more frequently they
widen from the base outwards and are divided by incisions of
variable depth into lobules. The flat surfaces of the cells are

occasionally furnished with spines or wart-like projections, and in

side or vertical view the cells are generally narrowly fusiform.
There is a central, plate-like chloroplast in each semicell, often
exhibiting irregular thickenings or ridges, and containing many
scattered pyrenoids. The zygospores are globose and furnished
with strong spines, simple or furcate at the apex.
There are 16 British species of the genus, none of which are really abundant,
although J/. denticulata Breb. (length 205 350 breadth 185 276 p.
JJL ; ;

fig. 61 C)
and M. truncata (Corda) Breb. (length 100138 ^ breadth 90 ;

129^; fig. 61 B) are widely distributed. N. rotata (Grev.) Ralfs (length

220240 n breadth 195220 M ) and J/. papillifera Breb. (length 135145 p
;
;
 166 Chlorophyceoe
breadth 115 145 /x)
are not uncommon
in permanent boggy pools and lakes,
whilst M. mucronata (Dixon) Wille and M. Jenneri Ralfs
oscitans Ralfs var.
are frequently found in the Sphagnum-bogs of mountainous areas. J/. furcata
Ag., M. Cnix-Melitensis (Ehreub.) Hass. (fig. 61 A) and M. pinnatifida (Kiitz.)
Ralfs are amongst the rarest and most elegant species of the genus.

Fig. 61. A, Micrasteriax Crux-Melitensis (Ehrenb.) Hass., from Bowness, West-

moreland (
x 365). B, HI. truncuta (Corda) Breb., from Thursley Common,
Surrey ( x 200). C, zygospore of M. denticulata Breb., from Halgavor Moor,
Cornwall ( x 110).

Genus Cosmarium Corda, 1834. [Dysphinctium Nag., 1849 ;

Kirchn., 1878
Calocylindrus (Nag.) Cosmaridium Gay, 1884
; ;

Pleurotceniopsis (Lund.) Lagerh., 1887.] This is the largest genus

of Desmids, embracing several hundreds of species, which although

exhibiting many varied characters, all conform to a common type

of structure. The cells are relatively short and the length is rarely
more than one-and-a-half times the breadth. There is a median
constriction of variable depth, in some cases very slight, but in
others exceedingly deep. The semicells may be circular, elliptical,
semicircular, ovate, pyramidate, or cuneiform in general outline,
and the apex, which may be rounded or broadly truncate, possesses
no apical notch. The cell-wall may be smooth, scrobiculate,
granulate, verrucose, or clothed with minute spines, the orna-
mentation in almost all cases being arranged upon some definite
plan. There is frequently a protuberance of some considerable
 Desmidiacece 107

size in the middle of each face of the semicell, so that the vertical
view, which is elliptical in some species, may in others be furnished
on each side with a more or less prominent swelling.
In the majority of species there is one central chloroplast in
each semicell, possessing four somewhat curved longitudinal plates
and either one or two large pyrenoids. In a few species there are

Fig. 62. A, Cosmarium reniforme (Ealfs) Arch, (a form), from Wicken Fen, Cambs.
(
x 473). B, C. firanatum Breb., from Chippenham Fen, Cambs. ( x 473). C
and D, C. ciramituin var. subgranatum Nordst., from Hornsey Mere, E. Yorks.
( x 473). E G, C. Meneghinii Breb.; E and F, from Hornsey Mere, E. Yorks.;
G, zygospore of a form from Bowness, Westmoreland ( x 473). H, C. prce-
mtirsum Breb., from Carrantuohill, Kerry, Ireland ( x 473). I and J, C.
bioculatum Bre"b. ;
from Rosvvell Pits, Cambs.; J, zygospore from Puttenham
I,

Common, Surrey x 473).

(
K
and L, forms of C. Reynesii Reinsch; L shows
one stage of cell-division ( x 1170). M, C. isthinium West, from Harris, Outer
Hebrides ( x 473). N, C. pseudoconnatum Nordst., from Capel Curig, N. Wales
(
x 473). /, front view; s, side or lateral view; r, vertical view.

several (from four to eight) parietal chloroplasts in each semicell,

each containing one or more pyrenoids.
Some investigators of these plants have attempted to establish
Nageli's genus Dysphinctium (= Calocylindrus Kirchn.) in order
to include certain species which are best left in the old genus
168 Chlorophycece
Cosmarium. The genus Dysphinctium can never be recognized
in a sound system of classification, as opinion must always remain
divided even upon many of the commonest forms that different
authors have included in it. Its characters are too indefinite and
artificial to be of
any Similarly, Pleuro-
real systematic value.

tceniopsis can never be established as a genus, as it would be a

small, polyphyletic assemblage, including a few strangely incon-
gruous species which occupy places far apart in the evolutionary
scheme of the genus Cosmarium. It must be remembered that
the primitive chloroplast of the Desmids is an axile one and
that the parietal condition has been independently acquired in
Cosmarium by a few scattered members of the genus. I have
already stated that if the large genus Cosmarium is ultimately
1

split up, the sections will not have to be based upon single
characters alone, but upon combinations of characters of which the
external form will be the most paramount. Until more is known
concerning the distribution of species of Cosmarium, the genus is
best left in its entirety.
The zygospores may be globose, angular-globose, cubical, or
almost of any outward form they may be smooth, scrobiculate,
;

furnished with simple or furcate spines of variable length, or

adorned with conical papillae. In a few species, such as G. melano-
sporum Arch., the spore-wall becomes perfectly black.
Asthere are 250 British species of Cosmarium it is not easy to point out
the chief peculiarities of the genus. The largest species which occurs in
Britain, and also one of the rarest, is C. ovale Ralfs (length 182 188ju;
breadth 100107 /*).
The smallest is C. subretusdforme W. & G. S. West
(length 7 '8 breadth 6'2 6'5 p.). The commonest species found in the
8 p ;

ponds and ditches of lowland districts are C. Botrytis (Bory) Menegh., C. prce-
morsum Breb. (fig. 62 H), C. xnbcostatum Nordst., C. humile Gay, C. granattim
Breb. var. subgranatum Nordst. (fig. 62 C and D), C. abbreviatum Racib. and
several forms of C. Meneghinii Breb. In the bogs of moorland districts
C. CtK'tu-bita usually abundant, and in upland Sphagnum areas
Breb. is

C. Ralfsii Breb., C. Breb., C. subtumidum Nordst., and others,

pyramidatum
are fairly general. Some species, such as C. Holmiense Lund., C. unceps Lund.,
C. subspeciosum Nordst. and C. Pokornyanum (Grun.) W. & G. S. West, are

usually found on dripping rocks, and C. Dovrense Nordst., C. microsphinctum

Nordst., and several others, prefer wet calcareous rocks. C. pygnnt'um Arch,

sometimes occurs in myriads amongst the leaves of submerged Sphagnum,

Genus Xanthidium Ehrenb., 1834. The cells of this genus
are somewhat flattened as in most species of Cosmarium, and the
1
G. S. West, 'Alga-flora of Cambridgeshire,' Journ. Bot. 1899, p. 115.
 Desmidiacece 169

median constriction is
invariably deep. The semicells may be
elliptical, trapeziform,hexagonal or octagonal in outline, and the
angles are furnished with simple or furcate spines. The presence
of these spines constitutes the
primary distinction between Xanthi-
dium and Cosmarium, and they are usually arranged in one plane
or in two parallel planes. In the centre of each semicell is a
thickened, scrobiculate area, or a protuberance of variable size,
and this character is the sole distinguishing feature between
Xanthidium and Arthrodesmus. In a few species the spines are

Fig. 63. A, Xanthidium a r ma turn (Breb.) liabenh., from Sligachan, Skye ( x 305).
B, zygospore of A. antilopaum (Breb.) Kiitz., from Ballyuahiuch, Galway,
Ireland ( x 365).

reduced to small apiculations at the angles. The chloroplasts are

parietal in most of the species and are arranged in the form of
cushions, four or six in each semicell, each with a pyrenoid. In
some of the minute species there is a single, central chloroplast in
each semicell, furnished with one
pyrenoid.
The zygospores are globose and adorned with blunt papillae or
long slender spines, simple or furcate at their extremities.
The genus was divided by Lundell in 1871 l into two sub-genera;
sub-genus Holocanthum, in which the spines are entire; sub-genus
Schizacanthum, in which the spines are forked at the apex. These
1
Lundell in Nova Acta Eeg. Soc. Scient. Upsala, 1871, ser. 3, vol. viii, p. 74.
 170 Chlorophycece
were put forward as genera by Wille but upon insufficient 1
,

grounds. Even in the most typical species which would fall under
Schizacanthum, namely X. armatum, the spines are sometimes
simple and in some of the tropical species there are numerous
2
,

intermediate stages between simple and much-forked spines.

There are 14 British species, of which a few are general but not Abundant,
and the rest are very scarce. X. armatum (Breb.) Raberih. is a large and
handsome Desmid occurring in moderate quantity in the bogs of the hilly
districts of the British Isles; length with spines 137 200 /x ;
breadth with
spines 88 110^; fig. 63 A. X. antilopceum (Breb.) Kiitz. (fig. 63 B) and
X. cristatum Breb. are not uncommon in certain areas, and X. concinnum Arch.,
which is the smallest species of the genus (length 9 9 5 p. breadth without
-

the apiculations 9'5 10'5/x), sometimes occurs abundantly amongst Sphagnum.

Species of this genus are regular constituents of the freshwater plankton.

Genus Arthrodesmus Ehrenb., 1838. This genus is primarily
distinguished from Xanthidium by the absence of the protu-
berance or scrobiculated
area in the centre of the
semicells. As a general
rule the spines are fewer
in number than in Xan-

thidium, and they are all

disposed in one plane.

The median constriction
is
deep and the semi-
cells, which may be
elliptical, trapeziform,
or subhexagonal in

shape, are furnished at

the lateral angles with

A C, Arthrodesmus Incus (Breb. ) one or two spines of

Fig. 64.
Hass., from Harrop Tarn, Cumberland; C, zygo- variable size and
spore (x365). D, A. Incus var. Ralfsii W. &
G. S. West, from Capel Curig, N. Wales ( x 365). strength. In the verti-
E, A. Incus var. validus W. & G. S. West (a form), cal view the cells are
from near Glenties, Donegal, Ireland ( x 474).
F and G, A. octocornis Ehrenb.; F, from Llyn always elliptical and the
Idwal, N. Wales; G, zygospore from Puttenham
poles are furnished with
Common, Surrey ( x 474). H
J, A. bijidus Breb.
var. truncatus West; H, from Strensall Common, spines. There is a single
N. Yorks.; I and J, from Kestou Common, Kent central chloroplast in
(x474). /, front view; v, vertical view.
each semicell contain-
1
Wille in Engler & Prantl's Natiirl. Pflanzenfam. 1890, p. 11.
2
West in Journ. Liun. Soc. Bot. xxix, 1892, p. 164, t. xxii, f. 1.
 Desmidiacece 171

ing one or two pyrenoids. The zygospores are globular, and the
outer surface may be smooth or clothed with simple, conical or
subulate spines.

Of the eleven British species, A. Incus (Breb.) Hass. is the only one that is
general and abundant. It is a small species, frequently met with in the con-

jugating state, and is one of the most variable of Desmids; length without
spines 13 33 p.; breadth without spines 13 28 p. length of spines 5
;
32 /x ;

fig. 64 A C. A. convergens Ehrenb. and A. octocornis Ehrenb. (fig. 64 F and

G) are not uncommon, but the other species are all rare.

Genus Staurastrum Meyen, 1829. This is the second largest

genus of Desmids and contains some hundreds of species of very
varied character, the majority of which possess a deep median
constriction. The semicells are elliptical, semicircular, oblong, or
cyathiform in outline, and the vertical view is generally triangular,
but may be quadrangular or polygonal. The angles are sometimes
rounded, sometimes acute, or they may be produced into processes
of considerable length. In many cases the angles of one semicell
alternate with those of the other. The cell-wall is occasionally
smooth, but it is more frequently adorned with a variety of spines
or wart-like excrescences, usually In
symmetrically disposed.
those species in which the angles are produced into processes the
latter are generally furnished with two or three strong diverging

spines at their extremities.

There is usually one chloroplast in each semicell, consisting of
a central mass with a number of radiating plates, varying from five
to eight (usually six; two in each angle). One pyrenoid is
present
in each chloroplast. In a few species there are several chloroplasts
in each semicell disposed in the manner of parietal cushions, but
intermediate states between this condition and a central mass are
not infrequent. This indefinite character was utilized by Lundell 1
for the formation of a
sub-genus, Pleurenterium, which has recently
been proposed as a genus'2 The absurdity of such a genus is
.

realized on considering the few incongruous forms it would have to

include. It should be borne in mind that the parietal condition
of the chloroplasts has been arrived at quite
independently by a
few widely different species of the genus Staurastrum.
The zygospores are globose or angular, rarely winged, but more
commonly clothed with long spines, which are simple or furcate at
1
Lundell in Nov. Acta Reg. Soc. Sclent. Upsala, 1871, ser. 3, vol. viii, p. 72.
-
Wille in Engler & Prantl's Natiiii. Ptianzenfam. 1890, p. 11.
172 CMorophycece
their extremities, and often situated each at the
apex of a mamillate
or obtusely conical protuberance.

Fig. 65. A and B, Staurastrum anatinum Cooke & Wills, from Llyn-y-cwui-ffynon,
N. Wales (x473). C, St. pitnctuhitnni Breb., from Esher Common, Surrey
(x473). D, St. polytrirlmm Perty, from Galway, Ireland ( x 365). E, St.
eloiigatiim Barker, from Bhiconich, Sutherland (x473). F, St. brachiatum
Half's, from Down, Ireland ( x 473). G, zygospore of St. furciijeniin Breb.,
from Pilmoor, N. Yorks. (x473). /, front view; v, vertical view.

There are more than 160 species of this germs known to occur in the
British Islands, but few ofthem are abundant. The most frequent species in
low-lying districts are St. pygmceuiii Breb., St. punctulatum Breb. (fig. 65 C)
and St. hexacerum (Ehrenb.) Wittr. In moorland areas St. margaritaceum
(Ehrenb.) Menegh. is general. The most abundant of the spiny species is

St. teliferum Ralfs. One of the largest British species of the genus is
St. tumidum Breb. (length 112 132 ^; breadth 91 103 p.) and the smallest
is St. iotirmnn Wolle. Some species, such as St. capitulum Breb., St. pileolatum
 .

Desmidiacece 173

Bn'b., St. Kjellmanii Wille (fig. 51 E), St. acarides Nordst. and St. Arndlii
Boldt, arc
principally confined to mountainous regions ; others, such as
St. pelagicum W. & G. S. West, St. & G. S. West,
pseudopelagicwn W.
St.jaculiferum West, St. paradoxum Meyen var. longipes Nordst., St. brevi-
apiitum Breb. and forms of St. anatinum Cke. & Wills tfig. 65 A and B), are
abundant in the plankton of lakes. A few of the most beautiful species of
the genus, amongst which may be mentioned St. Ophiura Lund., St. Cerastes
Lund., St. Arctiscon (Ehrenb.) Lund., St. verticillatum Arch, and St. longi-
spinum (Bail.) Arch., are confined to the extreme western districts of Scotland,
Wales and Ireland, and are most abundant in the plankton of those areas.

Genus Cosmocladium Breb., 1856. The cells of this genus

some of the smooth species of Cosmarium,
are similar to those of
but the individual cells are united by relatively thin mucilaginous
threads into branched colonies. Sometimes the entire colony is

Fig. 66. A, Cosmocladium constrictum (Arch.) Josh., from Pilmoor, N. Yorks. (

x 475).
B, C. pulchdhun Breb., from near Tarbert, Harris, Outer Hebrides (
x 475).
C, zygospore of C. perismm Roy & Biss., from the Clova Mts., Forfar (
x 475).
D F, Oocardium stratum Nag., after LiitkeiniUler ( x 730).

enveloped in a mucilaginous mass of much less density than the

connecting threads. There is one chloroplast in each semieell
containing a single pyrenoid. The zygospores are globose and
smooth, or they may be lobed (as in C. perissitm Roy & Biss.).
174 Chlorophycece
There are genus all of which are very rare.
five British species of this

66 A) and C. pulchellum Breb. (fig. 66 B) are

C. constrictum (Arch.) Josh. (fig.

perhaps more often observed than the others. All the species are very small,
the largest being C. Saxomcum De Bary; length 15 17 /x; breadth 13'5
14-5 /*.

Genus Oocardium Nag., 1849. This is the most extra-

ordinary of all the
genera of Desmids and usually occurs in large
colonies. The cells are small, slightly constricted and much
depressed, being considerably broader than their length. The
semicells are unequally depressed on the two sides, so that the

plant is symmetrical in one plane only. The vertical view is

broadly elliptical. There is one chloroplast in each semicell, con-

sisting of somewhat irregular plates radiating from a central mass

containing one pyrenoid.
The colony is generally hemispherical in shape, 1 2 mm. in
diameter, and occurs attached to calcareous rocks, not unfrequently
being itself encrusted with calcium carbonate. It consists of a
number of more or less parallel, radiating strands of mucus of
considerable thickness, each widening out towards the
strand
surface of the colony and occasionally branching. In the free end
of each mucous strand is lodged a single cell, disposed with its

longitudinal axis at right angles to the axis of the mucous strand.

The zygospores are unknown.
The only known species is Oocardium stratum Nag. It is extremely rare
and have only observed it in the limestone districts of West Yorkshire,
I

attached to rocks and stones in the beds of several mountain streams. Length
1316-5 p; breadth 18 19'5 M fig. 66 D F.
;

Genus Sphaerozosma Corda, 1835. The cells are small and

attached to form long filamentous colonies, often twisted and
sometimes enveloped in a mucous investment. The median con-
striction may be deep and narrow or it may be widely open, and
the semicells may be elliptical, oblong, or subrectangular in form.
The attachment of the cells is apical and is effected by small
rounded tubercles or short capitate processes. The vertical view
is elliptical. There is one axile chloroplast in each semicell,
furnished with a single pyrenoid. The zygospores, which are
globose or oblong, are either smooth or furnished with subulate

spines.
There are only five British species of the genus and none of them is
abundant. 8. vertebratum Ralfs (length of cells 19 /*; breadth 21 24 p;
 Desmidiacece 175

fig. 67 C) is the largest, and S. excavatum Ralfs (fig. 67 D F) and S. granu-

lutum Roy & Biss. are the most widely distributed.

Genus Onychonema Wallich, 1860. The cells are small and

form simple filamentous
colonies. The median con-
deep and narrow,
striction is
and the semicells are ellip-
tical or reniform,sometimes
with strong lateral spines at
each side. There are two
capitate processes of con-
siderable length attached to
each apex and disposed asym-
metrically. The cells are
united into long flexible
filaments by the overlapping
of these processes over the

adjacent cells. There is one

axile chloroplast in each
semicell, with a single pyre-
noid. The zygospores are
Fig. 67. A, Spondijlosium pulchellum Arch.,
globose and furnished with from Glen Shee, Perthshire (x365). B, S.
simple spines. papillatum W. & G. S. West, from Skipwith
Common, E. Yorks. x 475).
( C, Sphcero-
zosnia vertebratum Ralfs, from near CrdwalT,
There are three British species
Cornwall (x475). D F, Sph. excavatum
of the genus, all of which are Ralfs D, from Llyn Idwal, N. Wales ( x 475)
; ;

distinctly rare. The one most E,zygospore from Puttenham Common, Surrey
( x 475) F, zygospore from New Forest, Hants.
generally observed is 0. ;
filiformis x 475). G H, Onychonema Nordstedtiana
(Ehrenb.) Roy & Biss. (length of
(

Turner, from Strensall Common, N. Yorks.

cells 14 15 M breadth 14-5
;
(G, x475; H, x 730).
16 x.

Genus Spondylosium Breb., 1844. [Leuronema Wallich, I860.]

The cells are small or of medium size and are united by their
apices to form filamentous colonies, occasionally twisted and often
enveloped in a copious mucus. The median constriction is usually
deep and linear and the semicells are of very variable form. The
apices are flat or concave and the cells are joined merely by the
close apposition of their
apices, this being the sole distinguishing
feature between Spondylosium and The vertical
Sphwrozosma.
view is
elliptical, triangular, or trilobed. The chloroplasts are as
in Sphcsrozosma, and the zygospores are globose and smooth.
176 Chlorophycece
There are seven species of the genus known to occur in the British Islands,
none of which is abundant. S. papiUatum W. & G. S. West (length of
cells 8 9'5 p. breadth 9'5
;
1O5 p. fig. 67 B) and S. pidchellum Arch, (length
;

12 5-
15 p.; breadth 11 12'5 p.; fig. 67 A) are the most widely distributed.

Genus Hyalotheca Ehrenb., 1841. The cells are more or less

cylindrical and are connected

by their broadly truncate
apices into filamentous colo-
nies. The median constric-
tion very slight and the
is

semicells are trapezoid, sub-

quadrate or oblong in form,

with straight or convex lateral
margins. The filaments are

usually twisted and always

enveloped in a thick coat of
mucus. There are frequently
several slight swellings at the
base of each semicell near the
constriction, causing the cir-
cular vertical view to possess
two or three nipple-like pro-
jections at equal intervals
round the margin. There is

68. A dissiliens
one chloroplast in each semi-
Fig. C, Hyalotheca
(Sm.) Breb. A
and B, from Capel Curig,
; cell, consisting of a central
N. Wales (x365); C and D, zygospores
from Galway, Ireland (x 365). E H, mass with a number of radiat-
H. neglecta Racib., from the New Forest, ing plates, and containing one
Hants'. (x47o); E G, shosving aplano-
pyrenoid. The zygospores are
spores (); H, zygospore (z).
globose and smooth, and
in

H. dissiliens the four empty semicells unite to form a cruciform

structure which surrounds each spore.
There are four British species of the genus, of which H. dissiliens (Sm.)
is general and often abundant; length 1525^; breadth
Breb. 21 33^;
fig. 68 A C. This Desmid is more frequently found with zygospores than
any other. H. mucosa (Dillw.) Ehrenb. is a much scarcer plant, although
widely distributed. H. imdulata Nordst. (length 13'5 17'5/x breadth 7'5
;

9 p.)
and H. neglecta Racib. (length
2834-5 p.; breadth 11 '5 13 p.; fig. 68
E H) are amongst the rarest of British Desmids.

Series oblique junction of the

b. The new and old halves of
the cell-wall (at the region of the isthmus) develops an internal
 Desmidiacece 177

girdle-like thickening which projects into the old semicells during

the earlier stages of division. The cells are united to form thread-
like colonies.
Genus Desmidium Ag., 1824. [Didymoprium Kiitz., 1843 ;

Aptogonum Ralfs, 1848.] The cells are united to form twisted

filamentous colonies, often enveloped in a wide mucous coat. The

Fig. 69. A, Desmidium Swartzii Ag., from near Preston, Lancashire ( x 365).
B, D. quadrat um Nordst., showing cell-division ( x 475). C, zygospore of
D. cylindricum Clrev., from Donegal, Ireland (x350). D, zygospores of
D. aptofjonum Breb. ( x 475). E and F, Gymnozyga moniliformis Ehrenb.,
from Bhiconieh, Sutherland (x475); F, showing cell-division. G, zygospores
of G. inoniliformis var. gracilescens Nordst. ( x 475).

median constriction is
moderately deep and the semicells are much
depressed, so that the cells are generally much broader than their
length. The attachment of the cells is either by the close

apposition of their flat apices or by the apposition of corresponding

truncate apical projections. In the latter case there is a space of
variable width visible between the actual apices of two adjacent
cells. In vertical view the cells may be triangular, quadrangular,
or elliptical with mamillate poles. There is a single central
chloroplast in each semicell containing as many pyrenoids as there
are angles in the vertical view, and there are two longitudinal

plates diverging from each pyrenoid into the angle. The zygo-
w. A. 12
178 Chloi -ophycea*

spores are ellipsoidal, smooth or furnished with somewhat flattened

or conical papillae.

There are six British species of the genus, none of which is abundant.
tig. 69 A) and D. cylin-
D. Swartzii Ag. (length 14 19 p. breadth 37 43
; //, ;

dricum Grev. (fig. 69 C) are the two most general species. D. quadratum
Nordst. (fig. 69 B) and D. graciliceps (Xordst.) Lagerh. are two species very
rarely met with in the British Isles.

Genus Gymnozyga Ehrenb., 1840.

[Bambusina Ktitz., 1845.]
The cells are cylindrical and united by their flat
or barrel-shaped
ends into slightly twisted filamentous colonies. There is a slight
median constriction and at the base of each semicell is a swelling
of variable size. The vertical view is circular, sometimes with
two opposite papilke. The cell-wall frequently possesses delicate
longitudinal grooves. There is one central chloroplast in each
semicell with six radiating longitudinal plates and one pyrenoid.
The zygospores are ellipsoidal and smooth.
The only British species is O. moniliformis Ehrenb. (length 25 30 p. ;

breadth 17 5 22'5/x; fig. 69 E), which is generally distributed in boggy

-

districts, particularly in elevated localities.

Order IX. PEOTOCOCCOIDE^.

This order includes a large number of green Algae which are
UK >stly unicellular in character. The cells are commonly aggregated
to form irregular colonies and are often embedded in a
loose

copious mucilage. In a few of the forms a small multicellular

expansion is developed, and in others a definite coenobium which is
sometimes coenocytic in character. In some the cells are normally
ciliated and the plants are either motile unicells or more or less

complex motile colonies.

The orderincludes the lowest and most primitive of the green
Algae, forms through which most of the other Chlorophycese have
been evolved along divergent lines.
They are wonderfully varied in character and are found in
almost possible situations.
all The cell-walls may be extremely
delicate or firm and thick, and there is often a great development
of the gelatinous pectose compounds. The number and disposition
of the chloroplasts vary greatly in the different genera, and
pyrenoids may or may not be present.
 Protonxroidece 179

Most of the Protococcoidese multiply by simple cell-division,

which takes place in one, two, three, or sometimes in many
directions of space.
Asexual reproduction is generally accomplished by the pro-
duction of one or many biciliated zoogonidia from a mother-cell,
and in some families this is the sole method of reproduction.
There are many different types of asexual spores found in the
genera of the Protococcoidea5. Some of these are akinetes and
others are aplanospores. One type, which is worthy of special
note, is the autospore. A number of autospores are generally

produced in a cell, each spore having on liberation the general

form and appearance of the mother-cell. In some of the coenobic
forms the plants are reproduced by the formation of autocolonies.
Most of the methods of sexual reproduction, both isogamous and
heterogamous, which are exhibited by the various Chlorophycea?,
are found in the order Protococcoidese.
The order seems to me to be best subdivided into the following
eight families, of which the first one is doubtfully placed in the
Protococcoidese as the plants included in it have certain relation-

ships with the Chaetophorace*.

Family 1. Chfetopeltidece. Unicellular or multi cellular, .sometimes

pseudoparenchymatous. Some or all of the cells furnished with hairs
or bristles, either simple or sheathed and often mucous. Multiplication
by division of cells in two directions. Reproduction by 2- or 4-ciliated
zoogonidia and by 2-ciliated gametes.
Family 2. Volrocacece. Unicellular or consisting of a definite
co3nobium of cells, which are either united by protoplasmic processes
or enclosed within the swollen mucous mother-cell-wall. All the cells
are ciliated and motile in the vegetative state.

Family 3. Endosphwavece. Unicellular or slightly branched and

ccenocytic ;
cells solitary, generally rounded, often with button-like
excrescences of cellulose chloroplasts with numerous pyrenoids.
;
No
vegetative division. Reproduction by spores, zoogonidia and gametes.
All the genera are endophytic.
Family 4. Ckamciece. Unicellular cells solitary, differentiated
;

into base and apex, epiphytic on other Algee chloroplast parietal with
;

one pyrenoid. No vegetative division. Reproduction solely by zoo-

gonidia formed by successive divisions of the contents of a mother-cell.
Family 5. Pleurococcacece. Unicellular and globular, or of short,
ramified, few-celled filaments, never attenuated into hairs ;
often
pseudoparenchymatous chloroplasts one or several, parietal, with or
;

without pyrenoids. Multiplication by division in two or three directions,

and more rarely by zoogonidia. Cell-walls very firm.
122
180 Chlorophycece

Family 6. Hydrodictyacece. Thallus consisting of a cccnobium of

ccenocytes, non-motile, and formed by the apposition of quiescent zoo-
gonidia, which may or may not have escaped from the rnother-ca-uocyte.
Reproduction sometimes by resting-spores.
Family 7. Protococcacece (or Autosporacece). Cells solitary, free-

swimming, or colonial and associated in minute, more or less definite

colonies, easily dissociated or persistent. Multiplication by successive
division of contents forming autospores or autocolonies. Zoogonidia
rarely developed.
Family 8. Palmellaceoe. Mici'oscopic or macroscopic, gelatinous
and indefinite. Cells embedded
in a copious gelatinous envelope.

Multiplication by division in every direction cells often grouped in

;

twos or fours, sometimes with pseudocilia. Colonies free-floating or

attached. Zoogonidia with two cilia.

Family 1. CILffiTOPELTIDE,E.
The Algae included in this family consist of several genera of

very obscure affinities, all of which can be distinguished from other

members of the Protococcoidese by the presence of setse or bristles.
At present they known and the true nature of the
are but little
bristles has not yet been thoroughly worked out. They are uni-
cellular, or aggregates of loose cells, sometimes forming short
filaments or flat, pseudoparenchymatous expansions, which in some
instances appear to have arisen by a concrescence of short dicho-
tomous branch-systems.
The flat thallus of Chcetopeltis is similar to that found in the
Ulvacese, except from point of view of size and the fact that it is

attached by the whole of one surface. Chcetosphosridium is a

genus which may, perhaps, owing to the short creeping filaments
which it sometimes develops and the sheathed bristles, have some
relationship to the Herposteiracese, but its characters are so widely
different from those of Herposteiron that it is best kept
apart from
the Chsetophorales until our knowledge of the genus is augmented.
It may be that the resemblances between the Cha?topeltidege and
certain of the Cha?tophorales, such as they are, are due to a

parallelism of modification rather than to a direct affinity. This

has been illustrated in the table I have given of the phylogeny of
the green Algae on page 30.
In genera such as Chcetosphceridium and Conochcete multipli-
cation certainly takes place by the division of the .cells in two
directions.
 Chtftopelt idea 181

Reproduction has been observed to take place in several of the

genera by zoogonidia with two or four cilia, and by biciliated
isogamous gametes.
Throughout the entire family there is a marked dorsiventrality
such as appears elsewhere in the Coleocha3tacea3, Trentepohliaceae,
etc.

The following British genera are known :

A. Plants with a disciform thallus furnished with

scattered bristles Chietopeltis.
B. Plants consisting of a loose aggregate of cells in one stratum.
* Each cell with one seta or bristle.
Bristles with a basal sheath ; chloroplasts
parietal Ckoetosphceridium.
** Each cellwith several setce or bristles.
t Bristles few, with basal sheaths Conochcete.
tt Bristles many, without basal sheaths Polychcetophora.

Genus Chsetopeltis Berth., 1878. The thallus is a flat plate,

almost circular in outline, and consists of a single layer of compact

cells, more or less radiating from the centre. The growth of the
thallus is
peripheral as in Goleochcete, but the cell-walls are much
more gelatinous. From the upper surface of the thallus a number
of scattered mucous seta? arise, which are unseptate, simple, and
of considerable length. Each cell contains one parietal chloroplast,
which is often much lobed and perforated, and is furnished with a
single pyrenoid. Reproduction takes place by zoogonidia of which
2 8 arise from a mother-cell, each one possessing four cilia.
Isogamous gametes are also produced, similar in appearance to the

zoogonidia but with two cilia.

Ch. orbicularrs Berth, is not an uncommon plant in this country, occurring

as an epiphyte on the stems and leaves of various aquatic Phanerogams. The
thallus varies greatly in size and sometimes reaches a diameter of 1 mm.

Genus Chsetosphseridium Klebahn, 1892 The cells are 1

.

small and spherical, generally occurring in loose aggregates,

epiphytic on larger Alg* and other aquatic plants. They are

sometimes enveloped in mucus, but more often quite destitute of
a gelatinous investment. Klebahn states that the cells are com-
monly joined in a short filament by means of empty cylindrical
utricles, but I have not observed the presence of these structures

1
Klebahu in Jahrbiich. wissensch. Bot. xxiv, 1892, pp. 268282, pi. 4.
182 Chlorophyeece
in any British specimens. Each cell possesses at its upper pole a
small conical projection which forms a basal sheath for a long,
delicate seta or bristle. This
bristle extremely fine and
is

fragile, and is commonly broken

off at the apex of the basal
sheath. In most specimens it is

impossible to see any sheath-like

structure in the conical projec^-
tion, the bristle appearing to
arise from its apex. The chloro-

plast is
very variable, but in
some specimens it is
distinctly
parietal with one pyrenoid. The
division of the cells sometimes
takes place by the formation of
a horizontal division-plane, the
Fig. 70. AandB,
CJuetospharidium
globosum (Nordst.) Klebahn A, from lower
Esher Common, Surrey B, from Bow-
;

migrating daughter-cell
;

ness, Westmoreland. C, Ch. globosum

Asexual reproduc-
to the side.
var. depressum (West & G. S. West),tion occurs by zoogonidia, pro-
from the New Forest, Hants. (All
x 370.) duced four (or more ?) in a cell.
There is some little confusion
between this genus and one described by Borzi in 1892 as
Nordstedtia The latter was supposed to be founded on a plant
1
.

described by Nordstedt as Herposteiron globosa 2 and since referred ,

by Wolle to the genus Aphanochcete.' Klebahn, who compared

'

the original specimens of Nordstedt with Borzi's drawings of

Nordstedtia, affirmed that the latter represented an entirely
different plant. This being the case, one is compelled to accept
the genus Ckcvtosphwridium for the plants described by Nordstedt
"
as Herposteiron globosa."
Ch. globosum (Nordst.) Klebahn is widely distributed in the British
Islands, occurring chiefly in the Sphagnum-pools of permanent bogs. The
cells are 11 18 p in diameter, loosely associated, and occasionally surrounded
by a gelatinous envelope (tig. 70 A and B). There is a curious var. depress /ini
of this species with the cells much depressed (fig. 70 C).
The plant described as Cluetosphwidiiim Pringsheimii by Klebahn appears
to be identical with Aphanochoete globosa var. minor Hansgirg, and if it is
distinct from Ch. globomm it should be known as Ch. minus Hansg. Most
1
Borzi in Nuova Notarisia, iii, 1892, p. 50.
3
Nordstedt, Alg. aq. dulc. et Char. Sandvic. 1878, p. 23, t. ii, f. 22, 23.
 Chcetopeltidece 183

probably, however, it is but a small variety of Ch. globosum with the cells
9 12 p, in diameter 1.

Genus Conochaete Klebahn, 1893. The cells are comparatively

small, loosely aggregated, and embedded in a small amount of
mucus. Each cell is sub-
globose, often depressed,
and possesses a number
of delicate bristles which
radiate in all directions.
Each of the latter arises
either from the apex of
a mamillate protuberance
of the cell-wall or from
the base of an elongated
sheath. There are one or Fig. 71. A form of Conochcete comosa
two chloroplasts in each Klebahn, from the New Forest, Hants. (
x 370).
c/?, chloroplast ; o, oil-globule.
cell, each furnished with
one pyrenoid. The cells are markedly dorsiventral and sometimes
a prominent oil-globule is present in the basal half of the cell.
The plants multiply by cell-division in two directions and by
zoogonidia, of which four or eight are formed in each cell
2
.

Two British species are known, both of which are exceedingly rare.
C. comosa Klebahn possesses cells 13 26 p in diameter and the bristles,
which are 3 5 in number, are sheathed at the base (fig. 71). C. polytricka
(Nordst.) Kleb. possesses cells 10 16 p in diameter and the bristles arise
from the apices of a number of mamillate protuberances of the cell-wall, the
latter exhibiting a well-marked stratification.

Genus Polychaetophora West & G. S. West, 1903. The

plants are unicellular or sometimes composed of short, loosely
connected filaments of six or eight cells. The cells are subglobose
or ovoid, and the cell- walls are exceedingly thick and lamellose.
From the outer layers of this lamellose cell-wall from 8 to 12 long
flexuose bristles are given off. The bristles are very delicate and
attenuated to fine points they are quite simple, without any trace
;

of a basal sheath or any basal swelling, and they radiate in every

direction. Sometimes the cell-wall is unequally developed, a large
stratified outgrowth having been developed on one side. Such a
cell has a stalked appearance which presents many points of
1
This small form is known from N. Yorkshire vide West ; & G. S. West in
Trans. Yorks. Nat. Union, 1900, vol. v, p. 22.
2
Schmiclle in Hedwigia, 1899, p. 162, t. vi, f. 1619.
184 Ch lorophycece

resemblance to the cells in Borzi's genus Hormotila. Each cell

contains a single chloroplast, sometimes distinctly parietal, but

frequently subcentral and filled

with highly refractive oil-globules.
The presence of pyrenoids has not been definitely determined.

Multiplication takes place by division of the cells in two directions.

Fig. 72. Polychtetophora lamellosa West & G. S. West, from Cirencester,

Gloucestershire ( x 370).

P. lamellosa West & G. S. West is the only species of the genus and has
been found in Gloucestershire amongst Tolypothrix pygmoea. Diam. of cells

19_35 ^ ;
thickness of cell-wall 2'8 10'5 p.; length of bristles 86183 /*

(fig. 72).

Family 2. VOLVOCACE^E.
The either unicellular or
plants contained in this family are
they consist of definite coenobia of cells. They are distinguished
from all other Protococcoidese by being ciliated and motile in their
vegetative condition. The coenobia consist of a definite group
of cells either united together by protoplasmic processes or super-

ficially arranged
within the swollen wall of a mother-cell. The
number of cells in any coenobium remains constant so long as the
individual exists, and they all arose by cell-division while the

plant was still an embryo within the wall of the original mother-
cell.

The cells are rounded, angular, or ovoidal in form, generally

with a narrower anterior end to which are attached two, or rarely

The protoplasm of the anterior region of the cells
cilia. is
four,
 VolvocacecB 185

hyaline in character and often contains

a prominent pigment spot
and two (sometimes more) contractile vacuoles. The pulsation of
these vacuoles is alternate. There is a single nucleus, usually
occupying a central position in the cell. One chloroplast is present
containing one or more pyrenoids, and although its form is very
variable, it is generally more or less confined to the broader
posterior end of the cell.
Multiplication takes place by the division of the contents of a
mother-cell into or 8 daughter-cells, and, as the latter are new
2, 4,
motile individuals, the vegetative division in this family is strictly
homologous with the asexual reproduction by zoogonidia which is
found in the other groups of the Chlorophycese. In all the repro-
ductive processes in this family the plane of the first cell-division
is
usually at right angles to the longitudinal axis of the cell.
Sexual reprod action occurs in most of the forms by the union
of isogamous planogametes formed in a similar manner to the

vegetative daughter-cells, but in greater numbers. In some of

the higher forms there is a marked heterogamy, sexual reproduc-
tion being brought about either by the union of heterogamous

planogametes or by the fusion of antherozoids and oospheres. In

the zygospores or oospores the pigment spots of the two gametes
disappear, but the chloroplasts remain distinct and the cell-wall
becomes cuticularized. Germination takes place after a period of
repose.
As one passes from the lower to the higher members of the
VolvocaceaB there is a more striking progressive evolution of forms
than is exhibited in any other family of Algse. Therea gradual
is

replacement of isogamy by heterogamy, ultimately reaching the

highest condition of oogamy, and associated with this is an increase
in the number of cells and size of the ccenobium, accompanied by
a differentiation of vegetative from reproductive cells.
The plants of this family are of great interest on account of
the fact that they are the connecting links between the lower
forms of the green Algas and the Flagellata, a Protozoan group
which exhibits a mixture of animal and vegetable characters, and
from which it is now generally recognized that the Chlorophycese
have been evolved. There can be no doubt, however, in the mind
of anyone who has carefully studied the Volvocacese, that their
characters are such as to clearly separate them from the Flagellata
and place them as a family of the green Algae.
186 Chlorophycece
Plants of this family sometimes occur in prodigious quantity
in stagnant water, giving it a pale-green colour and a somewhat
unpleasant odour. They are occasionally the cause of foulness of
drinking-water, imparting to it a distinct oily taste 1 .

Excluding the Polyblepharideee, which exhibit a mixture of Flagellate and

Volvocine characters, the British Volvocacese are divided into the three fol-
2
lowing sub-families :

Sub-family I. Chlamydomonadece. Unicellular globose or ovoidal, ;

with a distinct but thin cell-wall. Cilia (or flagella) two, rarely four.
With one chloroplast of very variable form, usually including a single
pyrenoid.
Sub-family II. Phacotece. Unicellular, with the cells as in the
Chlamydomonadese, but with a thick solid cell-wall which separates
into two halves on the escape of the daughter-cells.

Sub-family III. Volvocece. Motile coenobia of Chlamydomonadiue

cells,usually embedded in a common mucilaginous investment more ;

rarely united by protoplasmic processes. All the cells are capable of

reproducing the plant or there is a differentiation into vegetative and
reproductive cells. Vegetative multiplication by division of some or
allof the cells to form daughter comobia. Isogamous or heterogamous
sexual reproduction.

Sub-family I. CHLAMYDOMONADESE.
This division of the Volvocacese includes a number of unicellular
Algae which are spherical, ovoid, subcylindrical, or rarely fusiform
in shape, and are provided with a thin cell-wall and two, or more

rarely four, cilia. The chloroplast is of variable form, but is

typically cup-shaped and occupies the posterior region of the cell,
more or less surrounding a central mass of protoplasm which
contains the nucleus. There is usually one pyrenoid, and a lateral
pigment spot is also generally present in the anterior region of
the cell.

Reproduction takes place by the division of the contents of a

cell which has come to rest into 2, 4, or 8 daughter-cells, each
of which soon acquires the characters of the mother-cell. The
successive division-planes are at right angles to each other and
the daughter-cells are ciliated motile individuals. Sometimes the
vegetative cells assume a palmelloid condition, in which they
1
Whipple in Trans. Amer. Micr. Soc. xxi, 1900.
:
Tliis classification is after 'Die Gattung Chlamydomonas und ihre
Dill,
iitichsten Verwanclten.' Jahrbiicher fiir wissenschaftliche Botanik. Berlin, 1895,
Bel xxviii.
 Volvocaeete 187

become rounded and repose in a copious mass of

mucilage.
Akinetes are also known to occur.
Sexual reproduction occurs by the conjugation of isogamous
or heterogamous planogametes, which are smaller but otherwise
similar to the vegetative individuals. They arise as do the latter
by the division of the contents of a mother-cell, but as many as
64 may be produced in one cell.

A. Vegetative cells with 4 cilia Carteria.

B. Vegetative cells with 2 cilia.
* Cell- wall thin, closely adherent.
t Cells spherical, ovoid, or ellipsoid, rarely
fusiform ;
chloroplast definite, with one
pyrenoid Cklamydomonas.
tt Cells fusiform, 3 or more times longer than
the diameter chloroplast indefinite, with
;

t\v< or more C-h lorogonium.

>
pyrenoids
** Cell-wall
thin, outstanding and connected by
protoplasmic threads Sphcerella.

Genus Carteria Diesing, 1868. [Pithiscus Dang., 1888; Cor-

biera Dang., 1888.] The cells are spherical, ellipsoid, or cordiform,
with a bell-shaped chloroplast containing a single large pyrenoid.
There is a prominent pigment-spot towards the anterior end of
the and there are four cilia. The only distinction between
cell,
this genus and Chlamydomonas is the presence of four cilia instead
of two.

C. multiflis (Fresen.) Dill is a

fairly abundant species in small pools, more
particularly of rain-water. Diam. vegetative cells 1016
p. (fig. 73 G). A
Genus Chlamydomonas Ehrenb., 1833. The vegetative cells
are of variable size, spherical, ovoid,
oblong-ellipsoid, or pyriform,
and the anterior end is often slightly produced to form a small
beak or rostrum to which are attached two cilia. The cell-wall is
hyaline, often very indistinct, and is closely adherent to the cell-
body. The chloroplast is very variable in form, and is usually
furnished with a single pyrenoid (rarely
entirely without or with
several pyrenoids). There are two (rarely mote) contractile vacuoles
in the anterior region of the cell, and generally a pigment-spot.
The reproduction typically that described for the sub-family,
is

and the wall of the zygospore may be smooth or

asperulate.
.*-
There are about 29 known species of the genus, but the characters of some
of them are not clearly evident. Little work has been done at the British
188 Chloi'ophycece

species of this genus, but CM. Kleinii Schmidle (length of cell 28 32/*;
breadth 812
p; fig. 73 J and K), Chi. De Baryana Gorosch. (breadth 12
20 [i ;
fig. 73 H and I) and Ckl.
pulvisculus Ehrenb. (
= Chl.
-E/irenbergiiQoTosch. breadth ;

14 26 /x) are not uncommon

in ponds, ditches and rain-
pools.
Concerning Chlamydom o-
nas Blackmail and Tansley 1
write, "This genus holds a
unique position among the
Green Algae,and indeed among
the whole of the Green Plants.
It may be regarded as the
phylogenetic starting point of
the various lines of Chloro-
phyceous descent. The history
of these is a history of the
K
B^KKijjW intercalation of a vegetative
H
phase between two successive
Fig. 73. A
G, Carteria multifilis (Fresen.) motile
Dill, from Bradford, W. York*. A and B, (Chlamydonionadiue)
generations, these motile
vegetative cells; D, gametes; E, conjugating
gametes; F, zygospore. H
and I, Chlamydo- phases being retained for re-
moiKts De Baryana Gorosch., from St Just,
productive purposes as zoo-
Cornwall. J and K, Chi. Kleinii Schmidle,
from Uxbridge, Middlesex (All x 4.75). cv, con- spores and gametes in the ;

tractile vacuoles; n, nucleus; p, pyrenoid; oogamous types the male

zg, gamete; 2, zygospore. gamete alone remains motile,
and constitutes in the Arche-
goniate series the last remaining representative of the Chlamydoinonadine
cell.
"
The co-existence within the limits of an undoubtedly natural genus of
the most primitive form of gamogenesis (the conjugation of equal clothed
gametes) with a gamogenesis which has the essential characteristics of true
oogamy is also a feature of unique interest."

Genus Chlorogonium Ehrenb., 1830. The vegetative cells

are fusiform, three or more times longer than the diameter, with a
thin cell-wall closely adhering to the body. There are two cilia
attached to the anterior extremity and a number of contractile
vacuoles scattered through the protoplasm. The chloroplast is ill-

defined, spongy and anastomosing in character, and contains four

or five, or sometimes many pyrenoids. France describes the chloro-
plast as forming a regular or irregular annular band, which may
split into a single or double spiral. Reproduction takes place by
1
Blackmail & Tansley in The New Phytologist, 1902, vol. 1, pp. 23, 24.
 Volvocacece 189

zoogonidia, four of which arise in a

mother-cell by the double
transverse division of the contents. Sexual reproduction also
occurs by the fusion of isogamous or heterogamous gametes.

Chi. euchlorum Ehrenb. (length up to 50 /A; breadth 8 12^i) is apparently

a very scarce British Alga of which I have examined a few specimens from
stagnant ditches near the Lizard, Cornwall.
Cercidium elongatum Dang. (1888) would be more correctly placed as
another species of this genus.

Genus Sphaerella Sommerfeldt, 1824. [Hcematococcas Ag.,

1828; Chlamydococcus A. Br., 1849.] The
cells are ovoid and very similar to those of

Ghlamydomonas, but the cell-wall stands out

away from the cell-body, being connected to
it
by delicate strands of protoplasm. The
chloroplast is bell-shaped and contains one
or several pyrenoids. The genus is scarcely
separable from Ghlamydomonas, only differing
in the outstanding cell-wall and the absence
of contractile vacuoles, the zoogonidia being

exactly of the Chlamydomonadine type.

Sph. lacustris (Girod.) Wittr. [Chlamydococcus
Fig. 74. A H. Sph<e-
pluvialis (Flot.) A. Br.] is abundant all over the rella lacustris (Girocl.)
country in ditches, rain-pools and bog-pools. The Wittr., from Bradford,
cells, which frequently become brick-red in colour W. Yorks. (
x 475).

owing to the presence of hsematochromin, vary from

8 to 30 p. in diameter they often become encysted, and the reproduction is
;

by zoogonidia and isogamous gametes (fig. 74). The curious phenomenon

known as "Red Rain" ovyes its colour in a few instances to the presence of
this Alga. Sph. nivalis Sommerf., which can scarcely be specifically distin-
"
guished from Sph. lacustris, is the Red Snow" plant.
Wille 1 has recently attempted to show that Hcematococcus is the correct
name of this genus, but I fail to see the reason for such a change. The remarks
made by Hazen 2 on the nomenclature of this genus should also be consulted.

Sub-family II. PHACOTE.E.

This sub-family is only distinguished from the Chlamydomo-
nadese by the thick, solid cell-wall, which on the escape of the
daughter-cells during reproduction separates into two portions.
1
Wille, 'Algologische Notizen X,' Nyt Magazin f. Naturvidenskab, Bd 41,
H. 1, Kristiania, 1903.
'-
Hazen, 'The Life-History of Sphairella lacustris,'' Mem. Torr. Bot. Club, vi,
1899, p. 236.
190 Chlorophycece
The two component parts of the cell-wall are sometimes evident
in the ordinary vegetative condition. The reproduction is similar
to that of the Chlamydomonadese.
Genus Phacotus Perty, 1852. The cell-body is ovoid in form
and considerably flattened, so that when viewed from the side it is
relatively narrow and biconvex. The cell-wall is thick, dark in
colour, with a rough or rugulose exterior, and consists of two
valve-like pieces. The chloroplast is large and parietal, with one
or many pyrenoids. There is a clear space of some size at the
anterior end of the cell, and two long cilia are attached imme-

diately opposite. The reproduction takes place by the formation

of 2 8 zoogonidia in a vesicle which bursts apart the two halves
of the mother-cell-wall by a longitudinal split.
P. lenticular is (Ehrenb.) Stein is not uncommon in
stagnant water.
Another genus of Pteromonas Seligo (1886) of which
this sub-family is

there are seven or eight species known from continental Europe. No doubt
some of these occur in the lakes of the British Islands, but as yet there are
no records of them.

Sub-family III. VOLVOCE.E.

The plants of this sub-family are composed of a motile coeno-
bium of Chlamydomonadine cells, generally embedded in a
considerable mucous envelope, and sometimes connected by proto- ,

plasmic processes. Attached to each cell are two cilia which

proj ect through the mucous coat and give the coenobium a rotatory
motion by their combined movements. The number of cells
present in a single coenobium varies from four in one species of
Gonium to 22,000 in some forms of Volvox. In the lower forms
all the cells are
equal and capable of reproducing the plant, but
in the higher forms there is a differentiation amongst the cells of
the ccEnobium, some of which are purely vegetative, whereas others
are solely reproductive cells.

Reproduction in the lower forms takes place by the formation

of a daughter-coenobium from every cell of the mother-coenobium.
The daughter-coenobium is formed within the wall of the mother-
cell, which swells up and ultimately sets the young colony free.
In the higher forms only certain of the reproductive cells, often
termed parthenogonidia, give rise to daughter-coenobia.
Sexual reproduction occurs in the lower forms by the union
of isogamous or heterogamous planogametes which arise by the
 Volvocacece 191

division of all the cells of the coBnobium. In the higher forms

there is a further differentiation elements into
of the sexual
antherozoids and oospheres, each of which arise from special repro-
ductive termed respectively androgonidia and gynogonidia.
cells

Genus Gonium Miiller, 1773. [Indus. Tetragonium West &

G. S. West, 1896.] The ccenobium is flat and plate-like, consisting

Fig. 75. A, Gonium pectorale Miill., from Cambridge. B F, G. lacnstre nob.,

from Esher Common, Surrey. E and F show the formation of new colonies
(
x 475).

of four or sixteen ovoid or slightly

polygonal cells arranged in one
plane and enclosed in a common mucilaginous investment. The
cells are connected by protoplasmic processes, and the cilia all
arise from one surface of the plate-like colony. The
chloroplast is
bell-shaped, much hollowed, and contains a single pyrenoid. Two
contractile vacuoles are present in each cell.
Reproduction takes
place by zoogonidia, which .are formed four in each cell or by the
dissociation of the colony into individual motile cells.
Conjugation
of isogamous gametes results in a
zygospore with a smooth or
rough cell-wall. Multiplication also occurs by the development of
daughter-coanobia from each of the cells of the mother-coenobium.
G. pectorale Miill. consists of a flat, somewhat
quadrate colony of sixteen
cells,each of which is 711
fj.
in diameter and of which twelve are
peripheral
and four central (tig. 75 A). It is a frequent and striking Alga in many
stagnant ditches and ponds, (r. fttcxxtre nob. [= Tetragonium lacustreW.
102 Chlorophycew
& G. S. West ] possesses a colony of four ovoid cells, each of which is
1

11-519 p. in length and 7 '5 15'5 /* in breadth; fig. 75 B F. The anterior

extremity of each cell is somewhat protracted and the cilia are only vibratile
towards their extremities, the movements of the ccenobiurn being correspond-
ingly sluggish. It is a much rarer plant than G. pectorale and prefers the

open water of large ponds and lakes. It may possibly be identical with G.
sociale (Duj.) Warming, but it appears to differ in the grouping of the cells
and in the nature of the cilia.

Genus Stephanosphsera Cohn, 1852. The coenobium consists

normally of eight cells arranged in an equatorial zone within a

tough, spherical or ellipsoidal investment. Occasionally young
coenobia are observed which contain only one or two cells. The
cells are ovoidal, and in the vegetative condition they possess

several green or colourless protoplasmic processes. The -zone of

cells is situated rather towards the posterior pole of the coenobium
and the cilia, of which one pair
is attached to the anterior pole of

each cell, penetrate the mucous investment in the equatorial plane.

The chloroplasts are rather irregular and contain from one to
several pyrenoids. Multiplication occurs by the division of each
cell of the ccenobium, after having assumed a more or less globular
form, into four or eight daughter-cells, each group forming a new
coenobium. Sexual reproduction occurs by the fusion of isogamous
gametes, 8, 16, or 32 of which are formed from the division of
a single cell. The gametes are fusiform in shape, conjugating
'

laterally to form spherical zygozoospores

'
which soon become
quiescent and of a yellowish-brown colour.
St. pluvialis Cohn, which is the only species of the genus, is known to
occur both in England arid Ireland. It is one of the scarcest of the Volvo-
cacesc and is usually found in the rain-water which collects in the hollows of
rocks. The cells are 6 12 p in diameter and the colonies 26 60 /j, ;

fig. 76 K.

Genus Pandorina Bory, 1824. The coenobia are spherical or

subspherical and usually contain 16 cells closely packed
within the
mucous investment. Sometimes coenobia of 8, or rarely of 32,
cells are observed. The cells are pyramidal in shape and reach
almost to the centre of the spherical colony, the pressure of
contact often causing them become quite angular.
to Two widely
divergent cilia are attached to the broad end of each cell. Multi-

plication takes place by the

formation of 16 -celled coenobia from
each of the cells of the mother-coenobium. The daughter-coenobia
1
West & G. S. West in Journ. Eoy. Micr. Soc. 1896, p. 160, t. iii, f. 113.
 Volvocacece 193

often remain within the old mucous investment of the mother-

cosnobium for some time, and the 16-ciliated colonies present the
appearance of a composite coenobium.

Fig. 76. A H, Pandorina morum (Miill.) Bory; A, adult colony; B, group of

daughter-colonies within the swollen mother-cell-wall; from near Bradford,
W. Yorks. (x475). C H, formation of zygospore (z) and its development
(after Pringsheim). K, Stephanosphcera pluvialis Cohn, ordinary vegetative
colony (after Hieronymus, x 320). g, gamete.

Asexual reproduction is by zoogonidia of a precisely similar

nature to the cells of Chlamydomonas The zoogonidia arise by
1
.

the longitudinal division of the contents of the mother-cell, and

each one secretes in addition to its own membrane a gelatinous
investment which ultimately forms the common investment of the
colony. Schroder 2 has also described an asexual method of repro-
duction in which the zoogonidia loose their cilia, vacuoles and
pigment-spots, and form a new colony. The gametes arise by the
division of the cells into 16 or 32 parts, and they exhibit con-
siderable variability in size. Sexual reproduction occurs by the
fusion of a pair of gametes, sometimes by the fusion of a smaller
active (male) gamete with a larger and more sluggish (female)
1
Dangeard in Le Botaniste, vii, 1900.
-
B. Schroder in J. B. Schles. Gesellsch. Vaterl. Cult. 1898, Zool., hot. sect.

W. A. 13
194 Chlorophycece

gamete. The zygospores possess smooth cell-walls and the

germination is indirect.

P. morum (Mull.) Bory is abundant in ponds and ditches all over the
country. The cells are 8 15 /x in diameter and the colonies 20 42 p. (fig. 76
A H).

Genus Eudorina Ehrenb., 1832. \Eudorinella Lemmermann,

1900.] The coenobium is globose or subglobose, rarely ellipsoid,
and consists normally of 32 cells distantly arranged within the

Fig. 77. Eudorina elegans Ehrenb. A, adult colony ( x 475) B, young colony
;

formed by division of contents of mother-cell x 730), from Frizinghall,

(

W. Yorks. C E, development of antherozoid-clusters from mother-cell ;

F, antherozoids (after Goebel).

periphery of a copious mucous investment. The cells are globose,

with a single bell-shaped chloroplast containing one or more
pyrenoids. Small colonies of only eight or sixteen cells are some-
times met with. The cilia (one pair) attached to each cell are
parallel until they reach the outer surface of the investment, when
they widely diverge. Multiplication takes place as in Pandorina
all the cells of the coenobium to form daughter-
by the division of
ccenobia. Sexual reproduction occurs by the union of fusiform or
pear-shaped antherozoids, produced 64 in a cell, with oospheres
which are slightly larger than the vegetative cells. The oospores
have smooth cell-walls and germination is direct.
There seems no valid reason for the separation of Lemmer-
 Votvoeacece 195

mann's genus Eudorinella 1

. Chodat 2 also considers that the two
described species of Pleodorina Shaw :{
should be regarded merely
as forms of Eudorina elegans, but the marked differentiation which
exists between the vegetative and reproductive cells of Pleodorina
does not support his view.

Eudorina elegans Ehrenb. is widely distributed in ponds, ditches and lakes

all over the British Islands. The cells are 10 25 p, in diameter and the
colonies 40 150 /*; fig. 77. The ellipsoid colonies of this plant are sometimes
described as possessing several mucous marnillate projections at one pole, but
I do not find this character in British examples, the colonies of which are
invariably globose.

Genus Volvox (L., 1758) Ehrenb., 1830. The ccenobium is

globose, consisting of a large number of small cells (200 22,000)

arranged in a single peripheral layer within the mucous investment.
The ccenobium is a hollow sphere and the cells are connected by
protoplasmic threads of varying stoutness. There is a differentia-
tion of the cells, the vast majority of them being purely vegetative

(somatic), and the remainder either parthenogonidia, androgonidia,

or gynogonidia. Each cell possesses a distinct chloroplast, two or
more contractile vacuoles, and a pair of cilia. Asexual reproduction
occurs by the development of new colonies from the partheno-

gonidia, of which about 8 (1 to 16) are found in a single coenobium.

These parthenogenetically formed individuals become detached in
the hollow cavity of the mother-coenobium and are ultimately set
free on its death. Sexual reproduction takes place by the fusion
of an antherozoid with an oosphere. The antherozoids arise by
the division of few or androgonidia, which are similar in
many
appearance to the parthenogonidia. The divisions of the androgo-
nidia produce either a plate-like or a spherical mass of antherozoids,
each of which is a small fusiform body, much attenuated at the
anterior end and furnished with two cilia. The gynogonidia are
few in number, larger than the vegetative and each one
cells,

becomes an oosphere. The oospores are globular, either smooth or

and their germination is direct, after a period of repose.
substellate,
Both asexual and sexual colonies occur, and the latter, which some-
times possess parthenogonidia, may be either monoecious or

1
Lemmermann in Berichte Deutsch. Botan. Gesellsch. xviii, 1900, p. 307.
-
Chodat in Beitrage zur Kryptogamenflora der Schwciz, Bd i, Heft 3, Berlin,

1902, p. 152.
Shaw in Botan. Gazette, 1894, six, p. 279 Kofoid in Bull. Illinois State Lab.
:i
;

v, 1898, p. 273; also in Ann. Mag. Nat. Hist. ser. 7, vol. vi, July, 1900.

132
196 Clilorophycece

dioecious. The most thorough investigations of the life-history

of the genus have been made by Klein 1
.

This genus isthe culminating point in the evolution of motile

cosnobic forms, and although the individual cells are of the

Fig. 78. A, C, and D, Volvox aureus Ehrenb. A, monoecious sexual colony x 210), (

from Bawcliffe Common, W. Yorks. D, ripe oospore (x475); C, two anthero-

;

zoids (after Klein). B, ripe oospore of V. globator (L.) Ehrenb., from Preston,
Lancashire (x475). a, androgonidia an, antherozoid g, gynogonidia.
; ;

primitive Chlamydomonadine type, the entire organism, by reason

of the differentiation of its component cells, has attained along its
own line a state of evolution quite comparable with the structural
differentiation attained along other lines by the highest known
forms of the green Algte.
The Rotifer Notommata parasitica sometimes occurs as a
parasite within the hollow coenobia of Volvox, swimming about
within the central cavity and feeding on the green cells either of
the adult or of the parthenogenetic daughter-coenobia.
1
Klein 'Morphol. u. biol. Studien iiber die Gatt. Volvox,' Pringsh. Jahrbuch. f.

wissensch. Bot. 1889, Bd xx.

 JEndcfsphfcracece 197

Two well-known species of Volvox exist in the deeper ponds and ditches
of the low-lying areas of the British Islands. They are often associated with
Lemna, and prefer ponds which receive a plentiful .supply of rain-water. The
adult colonies of V. globator Ehrenb. are 680 800 ^ in diameter, the cells are
very numerous, the protoplasmic strands connecting them are of considerable
thickness, and the ripe oospores are substellate (fig. 78 B). V. aureus Ehrenb.

is usually smaller than the preceding species, the adult colonies being 200

500 p, in diameter fig. 78 A, C, and D the cells are less numerous, the proto-
; ;

plasmic threads connecting them are extremely delicate, and the walls of
the ripe oospores are smooth.

Family 3. ENDOSPILEJRACEJE.
This family includes a number of endophytic Alga3 in which
the plant-body is either a simple rounded cell or a slightly
branched coenocyte, occurring wedged between the epidermal cells
of aquatic or marsh-loving plants. The cells are somewhat
variable in form and the cell-wall is sometimes thick and lamellose,
often developing on one side a button-like excrescence of cellulose.

Reproduction takes place by the formation of zoogonidia or

planogametes, or both. In the best known genus, Chlorochytrium
Cohn, the contents of the endophytic cell become broken up by
successive divisions, in a manner similar to that which occurs in
Characium, into a large number of small zoogonidia or gametes.
These are liberated by the gelatinization of the inner layer of the
cell -wall, which becomes protruded as a large vesicle such as is

found in the Pediastrese and many of the Protococcacese. The

biciliated gametes are isogamous and on their fusion the motile
4-ciliated zygospore comes to rest on the epidermis of the host-

plant. Germination takes place without any period of rest, the

new endophytic cell forcing a protuberance either between two
cells of the epidermis or into a stoma.
Some of these endophytic cells become akinetes and in this
condition they pass the winter.
There is a complete absence of ordinary vegetative division.
Seven genera have been described, but some of these, are
doubtfully distinct.
The researches of Klebs have shown that Cohn and other
of these
previous investigators were wrong in regarding certain
plants as parasites, and that not merely can the endophyte live
quite independently of its host-plant, but that the latter receives
no injury beyond the little mechanical pressure exerted by the
198 CMorophycecs

growing endophyte. The anomalous habit of these Algae is

explained by the protection afforded by the intercellular spaces of
1
the host. Freeman has suggested that the condition of affairs
met with in Chlorochytrium lends itself to the development of
parasitism and that the allied genus Phyllobium is
progressing in
that direction.
Genus Chlorochytrium Cohn, 1874. The cells are generally

solitary, globose, ellipsoid, ovoid, or irregularly curved and lobed,

Fig. 79. A, Vegetative cell of Chlorochytrium Lemnce Cohn, from Frizinghall,

W. Yorks. ( x 475). B D, Centrospheera Facciolace Borzi B and C, from
;

near Senens, Cornwall (

x 475) ; D, showing escape of zoogonidia (after Borzi,
x 410).

and they contain a large parietal chloroplast furnished with many

pyrenoids. Both starch and oil are often found in the cells.
Zoogonidia are formed by successive bipartitions of .the cell-

contents, at by perpendicular, and afterwards by radially

first

disposed walls. Gametes arise in a similar manner, and conju-

gation takes place within the vesicle formed by the protrusion of

the inner layer of the gametangium. Some of the species of this
genus are endophytes in the leaves of Lemna, Mentha, Rumex,
Lychnis, Sphagnum, etc., and others are marine, occurring in the
thalli of various marine Algse.

Chi. Lemnce Cohn occurs frequently in the leaves of Lemna trisulca diam. ;

cell. 40 143 p. ; fig. 79 A. Chi. Knyanum Szymanski is found on the leaves

of Lemna minor and L. gibbu.
The genus Stomatochytrium Cunningham (1888) differs from Chlorochytrium
mainly in the absence of the vesicle which surrounds the gametes in the
latter genus. This feature can scarcely be regarded as a generic difference.

1
Freeman, 'Obs. on Chlorochytrium,' Minnesota Botan. Studies, vol. n,
part in, p. 198.
 Characiece 199

Genus Centrosphaera Borzi, 1883. The cells are globose or

shortly ellipsoid and generally occur aggregated in a more or
less diffuse stratum beneath colonies of various members of the

Oscillatoriacese. The cell-walls are thick and lamellose, and possess

a projecting lamellose button of cellulose. Reproduction takes
place by biciliated zoogonidia, which usually arise in large numbers
from the zoogonidangia. Gametes have not been observed.
C. Facciolace Borzi 1 is known from the south of England. Diam. of vege-
tative cells 26 42 p ; zoogonidangia up to 80 fj. ; zoogonidia 2 3 p.
in breadth ;

fig. 79 B D.
There is scarcely any distinction between Centrosphcera Borzi, Endosphcera
Klebs and Scotinosphcera Klebs 2 the vegetative cells being almost alike in all
,

three. In Endospkcera the reproduction is by gametes, zoogonidia not having

been observed. In Scotinosphcera the only observed reproduction is by zoo-
gonidia, which arise according to Klebs in a most extraordinary manner.

Genus Phyllobium Klebs, 1881.The plants of this genus are

endophytic cosnocytes which occur in the leaves of certain marsh-
loving plants. They send out branching tubes through the inter-
manner of Pliyllosiphon
cellular spaces of the Iwst-plant after the

amongst the Siphonese. These branched tubes may or may not be

septate, and projecting from the surface of the leaf are small,
bright-green swellings. The latter contain a number of radiating
chloroplasts, each with a pyrenoid. Biciliated macro- and micro-

gametes occur.
dimorphum Klebs, which occurs in the leaves of Ajuga, Lysimachia,
P.
has not been observed from this country, but I have recently obtained
etc.,
what may prove to be another species from N. Uist, Outer Hebrides, thickly
studding the leaves of damp Sphagnum, the whole tissue of the leaf being
permeated by the branched tubes of the Phyllobium.

Family 4. CHARACTER.
The plants of this family are unicellular and they generally
occur as epiphytes, either solitary or in clusters, on other larger
Algae. The vegetative cells are of very variable form, but in most
cases they are attenuated and slightly oblique. There is always a
distinct differentiation into base and apex. The base is often
drawn out into a stalk of variable length with a disc for purpose of
attachment, and the apex is
generally acuminate. There is a

1 '
Borzi, Studi Algolonici I,' Messiua, 1883.
2
Klebs in Botan. Centralbl. 1881, xxxix, p. 1621.
200 Chlorophycece

single parietal chloroplast in each cell, generally of considerable

extent and containing one pyrenoid.
There no vegetative division.
is

Reproduction takes place by the formation of numerous

zoogonidia in each cell. There is a repeated division of the cell-
contents, several transverse divisions occurring before the first

longitudinal division, and in a short time each portion loses its

angular character, becomes rounded off, and forms a biciliated
zoogonidium. The escape of the zoogonidia takes place either by
a lateral or a terminal aperture. The pyrenoid disappears (in
some species) during the formation of the zoogonidia, and reappears
on their germination. Each zoogonidium on coming to rest

develops into anew plant.

Genus Characium A. Br., 1849. [Hydrocytium A. Br., 1855 ;

Hydrianum Rabenh., 1868.] The cells are fixed by a basal stalk,

Fig. 80. A and B, Characium Pringsheimii A. Br. A, from Mitcham Common, ;

Surrey; B, attached to a frustule of Tabellaria Jiocculosa, from Gunwen Moor,

Cornwall. C, Ch. subulatum A. Br., from Wimpole Park, Cambridgeshire.
D, Ch. ensiforme Herm., from Pilmoor, N. Yorks. (All x 520.)

usually rather short, but sometimes much elongated in form they ;

are spherical, ellipsoid, oblong, or fusiform, and they are generally

asymmetrical. The cells as a rule give rise to 16 or 32

zoogonidia.
Some eight or ten species of the genus are known to occur in Britain, of
which Ch. Sieboldii A. Br. (length 40 70 p. ;
diam. 20 33 /*), Ch. ambiguum
A. Br. (length 1530^; diam. 2'5 4 /x), Ch. Pringsheimii A. Br. (length
18 35 p.; diam. 5 ITS p; fig. 80 A and B) and Ch. ornithocephalum A. Br.
(length 19 33 p.; diam. 8 12*5 /*) are the most general. Ch. ensiforme Herm.
(length 65 86 /* ;
diam. 2'5 3'8 p ; fig. 80 D) is the most elongate species of
the genus.
 Pleurococcacece 201

Family 5. PLEUROCOCCACE.E3.
The plants
of this family are either unicellular or composed of

very short, slightly branched filaments, which consist of few cells

and are never attenuated to form hairs. The short filaments
are generally creeping and are often compacted into pseudo"-
parenchymatous masses. Considerable polymorphism is exhibited
by the different members of this family.
The cells are variable in outward form and contain one or
several parietal chloroplasts, with or without
pyrenoids.
Multiplication takes place by division in two or three directions,
and asexual reproduction occasionally occurs by the formation of
biciliated zoogonidia. Gametes are rarely produced.
In the two most frequent genera of the family, namely Plenro-
coccus and Trochiscia, the cells are more or less
globose and occur
aggregated in large masses the former prefers a subaerial habitat
;

whereas the latter prefers an aquatic existence. In Hormotila

there is a marked increase in thickness of the cell-wall at one side

only, branched colonies of cells being formed by the more or less

complete fusion of these remarkable lamellose excrescences.
Protoderma usually epiphytic, and its short cell-filaments
is

commonly assume a pseudoparenchymatous character it is ;

placed here on account of its close resemblance to certain of the

more uncommon states of Pleurococcus.
The cell-walls are usually very strong and firm, and the cells
are associated to form indefinite colonies or irregular aggregations.
This fact, and the complete absence of autospores, separate the
Pleurococcacese from the Protococcaceae.
Chodat includes in this family the genera Microthamnion and
1

Gongrosira, Alga? which I have referred to a distinct family the

Microthamniacese of the Chastophorales. There is considerable
resemblance between certain states of Pleurococcus and the young,
germinating plants of Microthamnion, but the latter genus has
reached a much higher stage of development than is ever attained
by forms of Pleurococcus. It is very probable that the family
MicrothamniaceaB has had a direct origin from certain of the
Pleurococcaceaa.

1
Chodat in Beitriige zur Kryptogamenflora der Schweiz, Bd i, Heft 3, Berlin,
1902.
202 Ghlorophycece
Wille 1 has placed the genus Trochiscia in the Volvocaceae in
close proximity to Chlamydomonas, but a careful consideration of
the facts of the case shows this change to be unjustifiable. The
ordinary vegetative condition of Trochiscia is a non-motile resting
state, zoogonidia rarely being produced whereas the normal ;

vegetative condition of Ghla/mydomonas (and indeed, of all the

Yolvocacese) is a motile one.
Genus Pleurococcus Menegh., 1842. [Protococcus Ag., 1824
(in part) ; Cystococcus Nag., 1849 ;
Chlorococcum Fries, 1825 (in
part); Cldorosphcem Klebs, 1883 (in part); Psendoplenrococcus
Snow, 1899.] The cells
are more or less globose,
sometimes angular by
compression, and they
occur frequently in

groups of two or four

due to imperfect separa-
tion after division. As
division occurs in three
directions a small cubic-
al colony is
occasionally
produced, but this easily
dissociates into its indi-
vidual cells. The cell-
walls are strong and
firm. There is one
Fig. 81, Pleurococcus vulgarix Meuegh.,
A, in
parietal chloroplast
from Cirencester, Gloucestershire. B, PI. ru-
fescens (Ktitz.) Breb. var. sanyuinens W. & G. S.
each cell, extremely vari-
West, from near Arncliffe, W. Yorks. (All x 520.) able in size and form,
rhJ, chloi'oplast; p, protoderma stage; pa, palmel-
loid state ; pg, bright red pigment ; pij, pyrenoid.
and with or without a
pyrenoid.
In moist places short filaments of cells are sometimes produced
which exhibit a simple type of branching they frequently radiate ;

from a few central cells of an angular, more or less parenchymatous

form. This condition is readily produced in cultures and can be
'
described as the Protoderma-stage.'
Vegetative multiplication takes place by division and sub-
sequent separation of the cells. Reproduction is brought about
1 '
Wille, Alg. Not. VII,' Nyt Magazin f. Naturvidenskab. 1901, F-istiania, B. 39,
H. 1, p. 9.
 Pleurococcacew 203

by the formation of one or many spores (aplanospores), by the

rejuvenescence of the contents of a mother-cell, by biciliated

zoogonidia, and by isogamous gametes. The polymorphism
exhibited by plants of this genus under various conditions of
humidity, temperature, etc., has caused much confusion with
regard to the identity of the different forms.
PL vulgaris Menegh. is one of the commonest of Algse, occurring in great
profusion in all kinds of damp situations, on stones,
walls, palings, tree-trunks,
etc., and usually forms a thin green incrustation on the windward side of
it

the objects on which it grows. The cells are as described for the genus and
the chloroplast is a massive lobed plate containing a prominent pyrenoid.
Diam. cells 9 20 p fig. 81 A. In PI. rufescens (Kiitz.) Breb. the cell-contents
;

are of a brick-red colour due to the presence of hpernatochroinin, which usually

appears to be dissolved in oily material. This species has a preference for
calcareous rocks a variety of it (var. sanguineus West & G. S. West) has
;

been observed in the limestone districts of West Yorkshire, forming large

brilliant blood-red patches,
covering those stones and rocks in the beds of
streams which could not be displaced by the rapidity of the current and
which are often left dry. Diam. cells 1120 p ; fig. 81 B.

Genus Trochiscia Kiitz., 1845. [Acanthococcus Lagerh., 1883 ;

Glochiococcus De Toni, 1888.] This genus is very closely related

Fig. 82. A F, Trochiscia aspera (Eeinach) Hansg., from Tremethick, Cornwall ;

A and B, vegetative cells C and D, formation of zoogonidia E, empty cell

; ;

from which zoogonidia (zg) have escaped F, palmelloid state.

; G and H,
T. hirta (Beinsch) Hansg., from Cambridge. I and J, T. panci.ipinosa West, from
Ben Lawers, Perthshire. K, T. reticularis (Beiusch) Hansg., from Keston
Common, Kent. (All x 520.)

to Pleurococcus, differing mainly in the external ornamentation of

the cell-wall and in the aquatic habit. The cells are perfectly
globose and usually occur in large aggregates in quiet water, or
rarely on damp ground. The cell-wall is either areolated or
thickly clouhed with denticulations, spines, or other prominent
204 Chlorophycece

processes. There are usually several parietal chloroplasts in each

cell, some which contain a single pyrenoid. Multiplication
or all of

rarely occurs by cell-division as in Pleurococcus, but reproduction

commonly takes place as in that genus by the formation of non-motile
gonidia or spores. Zoogonidia are rarely developed and the plants
sometimes pass into a palmelloid condition (vide fig. 82 F).
There are eight or nine British species of this genus, distinguished by their
external ornamentation. T. aspera (Reinsch) Hansg. (diam. veg. cells 13'5 29/* ;

fig. 82 A F), T. aciculifera (Lagerh.) Hansg. and T. reticularis (Reinsch)

Hansg. (diam. veg. cells 24 32 /* fig. 82 K) are amongst the most frequent
;

speciesmet with in quiet waters. T. hirta (Reinsch) Hansg. (diam. veg. cells
/i; fig. 82 G and H)
17 27 is often found on damp ground near the base of

tree-trunks.

[Westella De Wild.
2
Genus Radiococcus Schmidle, 1902 1 .
,

1897 (in part).] The plants consist of microscopic families of few

cells (generally four) arranged in a tetrahedral manner and

enveloped in a mass of jelly. The cells are rounded or occasionally

angular by mutual pressure, with firm cell-walls, and they contain
a single parietal chloroplast with one pyrenoid. Reproduction
takes place by the formation of four daughter-cells (spores)

tetrahedrally disposed within the wall of the mother-cell, which

ruptures and sets them free, the remains of the mother-cell-wall
persisting after the manner of Tetracoccus and Schizochlamys.
The genus distinguished from Tetracoccus by the smaller
is

families, which are enveloped in jelly, and by the tetrahedral

disposition of the cells. The plants often occur attached to the
under surfaces of water-lily leaves.
There are two species of this genus 3 of which R. nimbatus (De Wild.)
,

Schmidle ( = Pleurococcus nimbatus De Wild. Tetracoccus nimbatus Schmidle

; ;

Westella nimbatus De Wild.) is known from several parts of England. Diam.

of cells 8 15 p. In 1894 Schmidle 4 erroneously placed this Alga under
Ti'tracoccus and De Wildeman has further complicated matters by creating
the useless name Westella.

Genus Protoderma Klitz., 1843 ; Borzi, 1895. This genus

occupies a position in the Pleurococcacese by reason of the resem-
blance between it and certain states of Pleurococcus vulgaris. The
plants generally consist of a minute thallus of short cell -filaments
radiating from a small central group of pseudoparenchymatous
1
Schmidle in Allgem. Botan. Zeitschr. 1902, p. 41.
2
De Wildeman in Bull, de 1'Herb. Boissier, 1897, p. 503.
3
Schmidle in Hedwigia, 1902, Bd xli, Heft 4, p. 159.
4
Schmidle in Flora, 1894, Heft 1, p. 45.
 PleurococcacecB 205

cells. The cells are of a variable shape and the branches are
sometimes a little at-
tenuated. There is a
large parietal chloro-

plast in each cell con-

taining a single pyre-
noid.

P. viride Kiitz. (fig. 83

A C) is usually found
epiphytic on larger Algte,
such as Coleochcete orbicu-
laris, or on the stems and
leaves of aquatic Phanero-
gams, such as Callitriche,
Elodea, etc. I have pre-
1
viously suggested that
certain plants described
as Entocladia gradlis
Hansgirg
2
(
= Endoderma
gracileDe Toni) are most
probably referable to Pro-
toderma viride Kiitz. Fig. 83. A C, Protoderma viride Kiitz., from
Baildon, W.Yorks. epiphytic on Callitriche stag-
,

GenilS Hormotila nails; A'audB, outlines of colonies, x 520; C, single

x Hormotila mucigew Borzi, from
Borzi, 1883. Thevege-
& .
\ -

Boston Spa, W. Yorks. (x520).

tative cells are spheri-
cal, ovoid, or ellipsoid, rarely oblong, and from two to sixteen
of them occur within a more or less ample, firm, gelatinous in-

tegument, which is often concentrically lamellose. There is a

large chloroplast in each cell, frequently very granulose and destitute
of a pyrenoid. Multiplication takes place by cell-division, at first
in three directions,but subsequently in two, and finally in one
direction. In this way more or less moniliform series of cells are
*/

produced, all of which are connected by cylindrical lamellose

integuments. Zoogonidangia arise from vegetative cells by an
increase in and reproduction takes place by
size of the cell,

numerous minute zoogonidia, each with two cilia. Gametes have

not been observed.
The only known species is If. mucigena Borzi, an Alga which I have
observed from Boston Spa in West Yorkshire. It forms an expanded, dull

1
G. S. West in Journ. Bot. Febr. 1899, p. 58.
2
Hansgirg, Ueber Entocladia Eeinke und Pilinia Kiitz. ,' Flora, 1888, no. 33,
'

t. xii, f. 615.
206 Chlorophycece

green stratum on the damp surfaces of calcareous rocks. Diam. of veg. cells
4 12 /A zoogonidangia up to 30 p. in diameter zoogonidia 1
;
2'5 p in breadth
;

and 3 5 p. in length fig. 83 D. Chodat has observed Hormotila stages' of

;
i

a species of Pleurococcus, and he also suggests that they may be in part Dacty-
lotkece Braunii Lagerh., but with this latter suggestion I cannot agree.

Genus Urococcus Klitz., 1849. The cells are much as in

Hormotila Borzi, but are usually of a larger size and their contents
are coloured with a red-brown pigment. The cell-wall is thick
and lamellose, generally with a considerable increase in thickness
on one side. There is frequently an exuviation of the outer layers
of the cell-wall.
The genus
usually attributed to Hassall, but this is due to a
is

misconception. Hassall, in describing the characters of his " First

Subgenus" of Hcematococcus Ag., stated that "the term Ouracoccus

1

might be applied to the species of this subgenus." This suggestion

was improved upon by Kiitzing, who established the genus
Urococcus of Hassall but, if Urococcus is to remain as a genus, it
;

must be as " Urococcus Klitz. 2

'

U. insignis (Hass.) [
= Jfcematococcus insignis Mass.; Chroococcus
Kutz.
macrococcus Rabenh.] not uncommon in the bogs of moorland and upland
is

districts. Diam. of cells without integument 25 51 /x, with integument

41 78 p. It sometimes occurs in quantity amongst submerged Sphagnum,

especially in peaty pools.

Family 6. HYDRODICTYACEJE.
In this family of the Protococcoidea3 the plant-body consists of
a non-motile coenobium of coenocytes which floats freely in the
water. The coenocytes are arranged either as a flat plate or after
the manner
of a net, and they are of very variable form. In
Euastropsis there are only two cells (ccenocy tes ?
)
in the ccenobium,
in Pediastrum there may be more than fifty coenocytes, and in

Hydrodictyon there are often many hundreds, the ccenobium

reaching a length of several centimetres.
Multiplication sometimes occurs by the formation of autocolonies.
Reproduction takes place by the development of hypnospores
(usually aplanospores), by the formation of new coenobia by the
apposition of biciliated zoogonidia which have become quiescent,
and in some by the fusion of isogamous gametes to form a zygo-
spore. The germination of the zygospore is indirect.

1
Hassall, Bvit. Freshw. Alg. 1845,
p. 322. i,
-
Kiitziug, Spec. Algar. 1849, p. 206. Vide West & G. S. West in Journ. Bot.
June, 1897, p. 239.
 Hydroclk'tyacecK 207

The method of reproduction by the apposition of quiescent

zoogonidia to form new coenobia distinguishes the Hydrodictyacese
from all the other Protococcoidese, and the ccenocytic nature of the
ccenobium also distinguishes them from the coenobic forms of the
Protococcaceee (or Autosporacese).
It is most likely that Hydrodictyon and Pediastrum have no
direct affinity, the resemblance being due to convergence of
modification, but until more is known concerning the phylogenetic
relationships of these genera the Hydrodictyaceas are best divided
into the two following sub-families :

Sub-family I. Hydrodictyetx. New ccenobium formed by apposition

of zoogonidia within the mother-coenocyte. Plants macroscopic, coeno-
cytes arranged in the form of a net.

Sub-family II. Pediastrece. New ccenobiurn formed by apposition

of zoogonidia outside the rnother-ccenocyte. Plants microscopic, coeno-
cytes arranged to form a flat plate.

Sub-family I. HYDRODICTYE^.
This sub-family includes only the one genus Hydrodictyon.
The plants are macroscopic and consist of very large ccenocytes
which are disposed so as to form a more or less cylindrical net.
The zoogonidia swarm and become quiescent within the wall of
the mother-coenocyte, and there they become apposed to form the
new coenobium.
Genus Hydrodictyon Roth, The coenobium is a net-
1800.
and reaches a length of 8 10
like sac, freely floating in the water,
centimetres. The meshes of the net are of variable size and each
one is bounded by either five or six ccenocytes, the angles being
formed by the junction of three ccenocytes. The protoplasm of
each ccenocyte is confined to a lining layer containing many nuclei,
the central portion of the segment being occupied by a large
vacuole. There are no definite chloroplasts, the chlorophyll being
more or less diffuse through the whole protoplasm, but numerous

pyrenoids are present.

The normal method of reproduction is by the formation of a
very largenumber of zoogonidia within the mother-ccenocyte,
which swarm within the wall of the segment and then become
quiescent, immediately forming a reticulated daughter-ccenobimn
by the apposition of their extremities. The old cell- wall then
ruptures and the young ccenobium is set free. The zoogonidia are
208 Chlorophycece
biciliated they possess one nucleus and contain a single pyrenoid,
;

but as the young reticulum increases in size the pyrenoids of each

segment multiply rapidly. Timberlake states that the swarming
1

condition of the zoogonidia can be induced by the use of a reagent

composed of 100 c.c. of 1 per cent, solution of iridium chloride and
3 c.c. of glacial acetic acid. He points out that the cell-contents
first break up into large multinucleated masses, which in turn

Fig. 84. Hydrqdictyon reticulatum (L.) Lagerh., from the River Lea. A, nat. size ;

B, small portion of a young colony ( x 110) ; C, part of a large ccenocyte contain-

ing a very young colony (xllO); D, quiescent zoogonidia (x480); E, zoogo-
nidia which are becoming apposed to form a new colony ( x 480), p, pyrenoid ;

F, slightly older ccenocyte with four pyrenoids (p), x 480.

break up into smaller masses, until each mass contains a single

nucleus. Reproduction also occurs by the union of isogamous,
4-ciliated gametes, which escape from the mother-coenocyte by a
lateral pore. The gametes are smaller than the zoogonidia and
their escape is preceded by a swelling of an inner layer of the
cell-wall. This inner layer ruptures the outer layers and protrudes
as a vesicle in which the gametes swarm. On becoming free
they conjugate in pairs
2
. The zygospore is globose and after a
1
Timberlake in Botan. Gazette, xxxi, 1901, p. 203.
2
Klebs in Bot. Zeitung, xlix, 1891.
 Hydrodictyacece 209

short rest forms two or four large biciliated

zoospores, which on
coming to rest assume a polyhedral form. The repeated division
of the cell-contents of these polyhedral bodies results in the
formation of numerous zoogonidia, which by apposition give rise
to new net-like coenobia.
The division of the protoplasm of the adult cells of this genus
to form either zoogonidia or
gametes is a splendid example of free-
cell-formation.

The only known species, H. reticulatum (L.) Lagerh., which has received
the name of the " Water-net," is a very rare plant in Britain. The average
length of the adult ccenocytes just before they become zoogonidangia is 4 or
5 mm., but they are known to attain a length of 1 cm. The length of the
quiescent zoogonidia at the time of their apposition is 13'5 25 p.. The
swarming zoogonidia are 10 p in length by 8 p in breadth and the gametes are
a little smaller. Fig. 84.

Sub-family II. PEDIASTRE.E.

The plants of this sub-family are microscopic in size and consist
of a number of small coenocytes united to form a flat, disc-like
ccenobium. The zoogonidia swarm in a vesicle which is protruded
from the mother-coenocyte and the new coenobium is thus formed
outside the old coenocyte.
Genus Pediastrum Meyen, 1829. The coenobium is
always a
free-floating, plate, disc-shaped or star-shaped, and consists
flat

of a single layer of small coenocytes which is rarely duplicated in

the centre. The coenocytes are either parenchymatous and closely
united, or there are perforations of variable size between them
which give the coenobium a sieve-like aspect. The marginal
coenocytes are of different form from those in the centre and they
are generally furnished with a pair of diverging processes. There
is a single
chloroplast in each coenocyte, containing one pyrenoid.
The number of coenocytes in a coenobium varies from 2 to 64,
or even more. Coenobia of two coenocytes are very rarely observed,
and possibly belong to the genus Euastropsis, but in one species
(P. tetras) four is the regular number. The coenocytes are often
arranged in distinct rings round a central one, 8, 16, 32, or more
being the number in the coenobium. Nageli pointed out that
1

the coenobia were usually constructed as follows Colony of :

1
Nageli, Gatt. einzell. Alg. Zurich, 1849.

W. A. 14
210 Ghlorophycece

8=1+7; colony of 16 = 1 + 5 + 10 colony of 32 ;

=1+5+ 10 + 10 ;

but this arrangement is not always observed.

Multiplication occurs in some species by the formation of
autocolonies which arise by the division of the contents of a single

ccenocyte. Hypnospores are also frequently formed (fig. 85 E h).

Fig. 85. A, Pediastrum inteyrum Nag., from Ben Lawers, Perthshire ( x 475).
B, P. tricornutum Borge, from Glen Tuminel, Perthshire x 475). C and D,
(

P. tetrns (Ehrenb.) Kalfs, from Pilmoor, N. Yorks. (x475). E, P. duplex

Meyen, from Lough Fea, Londonderry, Ireland; hypnospores (h) (x475).
F H, P. Boryanum (Turp.) Menegh.; G, showing escape of zoogonidia;
H, young colony formed by apposition of quiescent zoogonidia F, from
;

Frizinghall, W. Yorks., x 475 G and H, x 220 (after Kerner).

; I, two marginal
cells of P. f/landuli fern in Benn., from Bisley Common, Surrey. J L, zoogo-
nidia and gametes of P. Boryanum (after Askenasy) J, zoogonidia and K,
;

gamete (x500); L, conjugation of gametes to form zygospores (z) (gametes

x 730, zygospores x 220).

Reproduction takes place by the successive division of the

contents of a coenocyte to form a number of zoogonidia, which are
suddenly liberated into an external vesicle through a slit in the
wall of the mother-co3nocyte. The zoogonidia swarm in this
vesicle and at length become quiescent, arranging themselves in
one plane as a new coenobium. Biciliated gametes have been
observed by Askenasy 1
,
which are much smaller than the zoogonidia
1
Askenasy, 'Entwickl. von Pediastrum,' Ber. Deutsch. Bot. Gesellsch. vi, 1888.
 Hydrodictyacece 211

and conjugate in pairs, the zygospore being polyhedral in form.

A new coanobium arises by the segmentation of the contents of
this polyhedral body.
The species of this genus exhibit great variation in the
characters both of their central and marginal cells, the length of
the marginal processes, etc. especially is this the case in cultures.
;

Their natural habitat ponds and ditches amongst other

is in small

water-plants, and not uncommonly in quiet bog-pools. Sometimes

they are numerous in the freshwater plankton.

The two most abundant species are P. Boryanum (Turp.) Menegh. (fig. 85
F H and J L) and P. M.r* (Ehrenb.) Ralfs (fig. 85 C and D), the former
sometimes reaching a relatively large size (200 p. in diameter). The marginal
processes of P. Boryanum are extremely variable and the cell-walls are often
granulated. P. tetras generally occurs in ccenobia of 4 (diam. 10'5 18 p.) or
8 (diam. 22 29 p.) cells. P. duplex Meyen 1829 (fig. 85 E) ( = P. pertusum
Kiitz. 1845) is also a widely distributed species. P. simplex Meyen (
= P.
A) are much rarer species.
clatliratum Lemm.) and P. integrmn Nag. (fig. 85
P. Boryanum and P. duplex are the most abundant species in the plankton.

Genus Euastropsis Lagerh., 1894 1

In this genus the
.

coenobium is
free-floating and consists of two flattened cells,

Fig. 80. Euatstropsis Richteri (Schmidle) Lagerh. A and B, from near Senens,
Cornwall (
x 760). C E, showing formation of young coenobia (after Lager-
heim ; highly magnified).

which are closely applied along their straight inner margins, the
outer margins being widely notched. The entire ccenobium has a
certain resemblance to a minute species of the genus Euastrum,
and was originally described as such-. There is a single parietal

chloroplast in containing one pyrenoid.

each cell
Lagerheim
describes the presence of one nucleus in each cell, but suggests
that there may be more. Reproduction occurs by oval, biciliated

Lagerh. in Tromso Museums Aarshefter 17, 1894, pp. 12

1
21, t. i, f. 8 27.
-
Schmidle in Flora, 1894, p. 60, t. 7, f. 25.
142
212 Chlorophycece

zoogonidia which swarm in a

as in Pediastrum.
vesicle On
becoming quiescent they arrange themselves in pairs and assume
the form of the adult cells, each pair forming one coenobium.

E. Richteri (Schmidle) Lagerh. is only known in the British Islands from

Cornwall. Length of coenobium 10 40 p., breadth 6 25 p, (fig. 86).

Family 7. PROTOCOCCACEJE (or AUTOSPORACE^3).

The Alga? included' in this family are free-floating, solitary
or colonial, and are most commonly associated to form minute
colonies of a definite construction. The
are sometimes firmly
cells
united to form a definite coenobium, as in Coelastrum and Sorasirum,
but as a rule the colonies easily dissociate into single cells or
smaller groups (or families) of
cells. With few exceptions there is
very mucus surrounding the colony. There is generally one
little

chloroplast (sometimes many) in each cell, parietal or occupying

the whole cell, and with or without a pyrenoid. There is one
cell-nucleus.

Multiplication takes place by the successive division of the

cell-contents either to form spore-like bodies which assume the
characters of the mother-cell before being liberated, or to form
colonies which on liberation resemble the mother-colony in all
except size. These are known respectively as autospores and
autocolonies. Multiplication by autospores, although such a
characteristic feature of the Protococcacea?, is not confined to this

family, as it occurs in Pediastrum, Radiococcus and certain other

genera.
Reproduction by zoogonidia or gametes is unknown except in
the genus Dictyosphasrium.
The Alga? of this family are well marked off from the Pleuro-
coccacea? and Palmellacese by their definite colonies and by their
method of multiplication. They are little removed from the most
primitive forms of green Alga? and a few of them are capable of
profound modification by cultivation in different media. In their
natural state, however, they exhibit a remarkable constancy of
character and many of them are ubiquitous in all climates.

The family is best subdivided into the seven following groups :

Sub-family I. Codnstrcce. Cells primarily globose or broadly

lunate, forming a definite spherical or polyhedral coenobium.

Sub-family II. Crucigeniew. Cells of variable form, arranged in a
regular flat plate.
 Protococcacece 213

Sub-family III. Sehma&trece. Cells elongated, ofteu greatly at-

tenuated and frequently curved; solitary or associated in definite or
loosely coherent colonies. One chloroplast in each cell.
Sub-family IV. Oocystidece. Cells globose, ellipsoid, reniform or
sometimes sublimate. Daughter-cells generally retained in the en-
larged wall of the mother-cell. Several or many chloroplasts (rarely
one) in each cell.
Sub-family V. Tctmedriece. Cells solitary ;
flattened and angular,
with a definite number of angles, or tetrahedral, octahedral or poly-
hedral. Angles generally furnished with simple or furcate spines.
Sub-family VI. Phytheliece. Cells globose or ellipsoid, solitary or
colonial, furnished with two or more long attenuated bristles.
Sub-family VII. Dictyosp/Keriece. Cells globose, ellipsoid or ovoid,
associated to form colonies, and joined more or less completely by the
persistent old walls of the mother-cells, which sometimes become trans-
formed into special connecting-threads.

Sub-family I. CCELASTRE^l.
This is the only group of the Protococcaceae in which there is a

definite and regular coenobium of spherical or polyhedral form.

The globose or polygonal, with or without short
cells are either

projecting processes, by means of which they are united to form a

hollow sphere or they are broadly lunate and united at the centre
;

of the spherical colony by short stalks. Multiplication is by the

formation of an autocolony in each cell of the coenobium, which is
ultimately set free by the dissolution or splitting of the mother-
cell-wall.
Genus Coelastrum Nag., 1849. [Hariotina Dang., 1889.] The
coenobium, which is hollow and is com-
is
spherical or polyhedral,

Fig. 87. A, Coelastrum cambricuin Archer, from Lough Gartan, Donegal, Ireland.
B D, C. sphcericum Nag. B and C, small coenobia from near Penzance,
;

Cornwall; D, large cceuobia giving rise to daughter-coenobia (autocolonies),

from Bowuess, Westmoreland. (All x 475.) '
 214 Chlorophycece

posed of a variable number of cells united by their lateral margins

to form a single peripheral layer. In some species the cells are
globose, in others more or less angular, and in others they are
furnished With projecting processes by means of which they are
joined together. The latter forms exhibit intercellular spaces of
variable size and the cells are often furnished with a truncate

projection (rarely two) on their free outer surfaces. The maximum

number of cells observed in a single colony is 64, but in most

species the number is 8, 16, or 32. Each cell contains a single

chloroplast with one pyrenoid. The multiplication is by typical
autocolonies, which are liberated by a split in the wall of the
mother-cells or more rarely by the entire gelatinization of the
mother-cell-walls. Single spores are sometimes developed from
individual cells.

The two most frequent species are C. sphterwum Nag. (diam. ccenob.
1892 p diam. cells 423 ^ fig. 87 B D) and C. cambric urn Arch.
; ;
1868
(
= C. pulchrum Schmidle 1892); fig. 87 A. In the former species the cells
aresomewhat conical with a polygonal base, and in the latter species they are
more or less distinctly lobed arid furnished at the same time with a truncate
surface projection. C. cubicum Nag., C. microporum Nag., C. verrucosum
Eeinsch and C. proboscideum Bohlin are species of considerable rarity.

Fig. 88. Ccelastrum reticulatum (Dang.) Serm, a small irregular form from
Churchill, Donegal, Ireland x 475).
(

C. reticulatum (Dang.) Senn, which was made the

type of the genus
Hariotina by Dangeard, differs from all other species in the nature and dis-
position of the processes of the cells. The ctenobium consists of 4, 8, or 16
cells,and the young autocolonies are retained for a relatively long period
within the walls of the mother-cells. The processes of attachment of the cells
are narrow prolongations, often curved, and sometimes irregularly
disposed.
This plant has also been named C. subpulchrum by Lagerheim and C. distans
by Turner, but Dangeard's name takes precedence.
1
It is known from
2
Donegal, Ireland and large
,
colonies are not infrequent in the plankton of
Lough Neagh. Diam. cells 6 24 p. fig. 88. ;

1
Dangeard, 'Memoire sur les Algues,' Le Botaniste, 1889; Chodat & Huber in
Bull. Soc. Bot. France, torn, xli, 1894.
2
West & G. S. West in Journ. Bot. March, 1903.
 Protococcacece 215

Genus Sorastrum Kiitz., 1845. [Selenosphcerium Cohn, 1879.]

The coenobium is globose, consisting commonly of 16 to 64 (rarely
4 or 8) stalked cells, the
stalks uniting in the
centre of the co3nobium
to form a small faceted
sphere. In small colonies
the stalks of the cells are
very short and the central
sphere is not always ap-
parent. The cells are
v -M i i .
Fig. 89. Sorastrum spmulosum Nag., A,
broadly sublimate, rein- small coenobium from Pilmoor,N.Yorks.(x 475);
form or subtriane'llHr in ^' ^ ai'8 e coenobium from Clifden, Galway, Ireland
(
x 450).
shape, and each extremity
is furnished with two
spines (rarely one) of moderate length.
There is one chloroplast, containing a pyrenoid, in each cell.
The multiplication by the formation of autocolonies, but the
is

details have not been worked out.

TheBritish species usually met with is S. spinulosum Nag. diam. of ;

ccenobia 31 90 fj.; length of cells without spines 11 26 //,; fig. 89. It is a

scarce plant occurring in bog-pools or amongst aquatic Phanerogams in the
quiet margins of lakes. S. Americanum (Bohlin) Schmidle occurs in the
freshwater plankton of the Outer Hebrides.

Sub-family II. CRUCIGENIE^E.

The ccenobia consist of few or cells regularly arranged in

many
the form of a flat
plate. The
are very variable in form,
cells

generally somewhat rounded, and occasionally furnished with spines.

They are disposed in groups of four, the latter being held in

position by a tough mucilage. In some the chloroplasts possess a

single pyrenoid, but in others pyrenoids are absent.
The multiplication is by autocolonies, which in some cases are
set free almost immediately, but in others remain for some time as
part of the mother-colony.
Genus Crucigenia Morren, 1830. [Stan'royenia Kiitz., 1849 ;

Lemmermannia Chodat, 1899 ;

Willea Schmidle,
1900.] The
coenobium consists of 4, 8, 16, or 32 cells arranged as a flat plate
and held in position by a mucilaginous envelope, but under
favourable conditions of environment as many as 128 cells have
210 Chlorophycece
been observed in one colony Even in the large colonies the cells
1
.

are distinctly arranged in groups of four, the cells of each group

being closely adherent except in the centre, where there is usually

a small quadrate or rhomboidal space. The cell-walls are smooth,
and each cell contains a single chloroplast with or without a
pyrenoid. Multiplication occurs by autocolonies of four colls pro-
duced by the cruciform division of the contents of a mother-cell.
Schmidle 2 has given a good account of this genus under the
name of Staurogenia,' and has also described
'
the formation of
hypnospores.
Chodat has separated the genus Lemmermannia from Crucigenia
mainly owing to the absence of a
pyrenoid from the chloroplast. For
the present I prefer to regard the two
as identical. There is no difference
in the structure of the colonies and
the mode of multiplication is pre-

cisely the same in each, the daughter-

cells persistently remaining as parts
H of the mother-colony in several

Fig. 90. A C, Crucigenia rect- species of Crucigenia as well as in

angular is (Nag.) Gay, from Lough Lemmermannia. In all the speci-
Shannacloontippen, Galway, Ire-
land; C, with formation of auto- mens I have observed of Crucigenia
D and E, C. quadrata
spores.
Morreu, from Settle, W. Yorks. F,
Tetrapedid, which was the plant upon
C. Tetrapedia (Kirchn.) W. & G. S. which Chodat founded the genus
West, from plankton of L. Neagh, Lemmermannia 3 there was the same
Ireland. G and H, Tetrustrinn ,

Staurogenieeforme (Schrod.) Chod.,

small gap in the centre of each group
from near Kievaulx Abbey, N. of four cells that is present in all
Yorks. (All x520.)
other species of Crucigenia.

C. rectangularis (Nag.) Gay is the most frequent species of the genus;

length of cells 5 9 p breadth 4 6 p fig. 90 A C.

;
C. quadrata Morren
;

1830 ( = ?(?. triangnlaris Chod. 1900) is a much rarer species; diam. cells
55-5 M; fig. 90 D and E. Tetrapedia (Kirchn.) W. & G. S. West
C. 4

(
= Lemmermannia emarginata Chod.) is known from the plankton of Lough
1
West & G.
S. West in Ann. Bot. xii, 1898, p. 36.
2
Schmidle in Berichte Deutsch. Bot. Gesellsch. 1900, Bd xviii, pp. 149157.
Under the heading of Lemmermannia emarginata, Chodat (in Beitriige zur
:i

Krypt.-fl. Schweiz, Bd I, Heft 3, 1902, p. 222) makes some irrelevant remarks

concerning Tetraedron pentaedrica W. & G. S. West (which, for some reason, he
seems to think was described as a Tetrapedia !) and Tetrapedia morsa W. & G. S.
West, which clearly show that he is quite unacquainted with either the published
descriptions or figures of the plants in question.
4
West & G. S. West in Trans. Eoy. Irish Acad. vol. xxxii, sect. B, part i, 1902, p. 62.
 ProtococcacecB 217

Neagh, Ireland ;
diam. cells 4'8 9'5 fj.;
diain. cu'iiob. of 4 cells 10'5 l~>~~>
p.;

fig. 90 F. All the species are regular constituents of the freshwater plankton.
C. irregularis Wille 1 is known from the plankton of several lochs in the

Shetlands, and from Norway. It is a most interesting species, differing from

C. rectangularis only in the somewhat irregular colonies and the absence of

pyrenoids from all the cells of the colony. Some of the cells in a colony of
C. rectangularis are often destitute of pyreuoids, and Wille was quite right
in placing his Norwegian plant (C. irregularis) as a species of Crucigenia.

Unfortunately, however, the useless generic name Willea has been put for-
' '

ward by Schmidle 2 to include Crucigenia irregularis. This genus is simply

founded upon the absence of pyrenoids from all the cells of the colony and ;

Lemmermann 3 has since placed Willea,' as a subgenus of Cohniella, also

'

owing to the absence of pyrenoids! Surely, in Willea, Lemmermannia, and

Lemmermann's suggestion that Willea should be a subgenus of Cohniella, the
climax of absurdity has been reached with regard to the presence or absence
of pyrenoids as a generic distinction. No one who has been fortunate enough
to observe colonies of Crucigenia. irregular!* could dispute their close affinity
with forms of C. rectangularis. The former species has most likely had a
direct origin from the latter.

Genus Tetrastrum Chodat, 1895. [Cohniella Schroder, 1897.]

The coenobium consists of four cells arranged in one plane and
surrounded by a thin mucous envelope. The cells are rounded or
somewhat angular, and they possess from two to five spines of
variable length on their external margins. The multiplication is
by the formation of autocolonies of four cells in each mother-cell.
Pyrenoids may be present or absent. The genus only differs from
Crucigenia in the presence of the marginal spines and in the
regular 4-celled condition of the coenobium.
The only two known species of the genus are T. heteracanthum (Nordst.)
Chod. and T. Staurogeniceforme (Schrod.) Chod. The former is known from
West Ireland and the latter (fig. 90 G and H diam. of cells without spines
;

3 6 p.) occurs in North Yorkshire.

Sub-family III. SELENASTRE.E.

This group of the Protococcacese is characterized by the elon-

gation of the cells, which are often very narrow with the extremities
attenuated to fine points. They are frequently lunate or arcuate,
and may be solitary or associated to form colonies of a more or less

Wille, Algologische Notizen IV,' Nyt Magaziii for Naturvideiiskb. Bd 38,

1 '

Heft 1, 1900, p. 10, t. 1, f. 15.

2
Schmidle, 1. c. p. 157.
3
Lemmermanu in Berichte Deutsch. Bot. Gesellsch. 1904, Bd xxii, p. 22.
218 ChlorophycecB

fragile character, the cells in some instances being held in position

only by the enveloping mucus. There is a single chloroplast in

each cell, which may contain one or several pyrenoids or may be
destitute of them. In rare instances the chloroplast is fragmented.
The cell-wall is firm but delicate.

Multiplication principally by autospores and autocolonies,

is

and this often takes place by the oblique division of the contents
of the mother-cell.

The following genera of the Selenastrese are British :

A. Cells or colonies almost destitute of mucus.

* Cells attenuated to acute
apices multiplication by
;

oblique division of contents of mother-cell, the

daughter-cells often remaining loosely attached
by their apices Dactylococcus.
** Cells
ellipsoid or much attenuated, forming more
or less definite colonies consisting of a row of
cells in one plane Scenedesmus.
*** Cells sublimate or
ellipsoid, arranged in groups
of four in one plane, the groups being attached
to form an irregular colony Dimarphococcus.
**** Cells
elongate and acutely attenuated, often
lunate, solitary or loosely grouped in irregular
bundles.
moderate length, usually with not more
+ Cells of
than one pyrenoid Ankistrodesmus.
ft Cells greatly elongated, pyrenoids numerous... Closteriopsis.
***** Cells or often at-
ovoid, oblong, club-shaped,
tached by their apices to form radiating colonies Actinastrum.
****** ( e ii,s attenuated,
lunate, arranged back to
i

back to form definite colonies Selenastrum.

B. Colonies enveloped in a copious mucus.
Cells lunate or much curved, disposed more or less

irregularly in the enveloping jelly Kirchneriella.

Genus Dactylococcus [Indus.

Nag., 1849.
Coccomyxa,
Schmidle, 1901.] The cells are ellipsoidal, fusiform or pyriform,
often sublimate and generally with acute apices, which are some-
times unequally prolonged into spine-like processes. They occu'
solitary or loosely connected by their acute extremities to fori
fragile colonies of few cells. The chloroplast is single and pariet;
and sometimes contains a single pyrenoid occasionally it becomes
;

fragmented into two, three, or four parietal pieces. Some species

are truly aquatic, but most of them occur on wet rocks and moist
surfaces, forming a dark green mucous stratum. Multiplication is
 ProtococcaeecB

by the oblique longitudinal division of the contents of the mother-

cell.

D. bicaudatus A. Br. is a lunate species with the apices greatly prolonged

and its chloroplast is destitute of a pyrenoid length of cells 13

;
39 p diain.
;

of cells 2-5 5-8 /u; fig. 91 A. D. dispar W. & G. S. West is unequally

Fig. 91. A, Dactylococcits bicaitdutus A. Br., from near Bradford, W. Yorks.

B and C, D. bicamlatus var. subramotiiis W. & G. S. West, from Witldale Fell,
N. Yorks. D, D. dispar W. & G. S. West, from Dorking, Surrey. (All x 520.)

developed at the extremities and the cells frequently become irregular;

fig. 91 D. The Alga described as D. Debaryanus Reinsch, which often occurs
in large numbers as an epiphyte on Cyclops and other small Crustacea, is a

species of Characium.
There is little doubt that D. infusionum Nag. is merely a state in the life-

history of Scenedesmus obliquus (Turp.) Kiitz. It is usually aquatic and often

occurs in immense quantity in the water of flower-pots and in aquaria. It
exhibits great variability of form and its chloroplast commonly contains a
pyrenoid length of cells 7'5 19 /*, breadth 2'8 5 8 p.
-

Genus Scenedesmus Meyen, 1829. In this genus there is a

coenobium of more or less definite form, consisting of a variable
number of cells arranged either in a single row or in two alterna-

ting rows. The cells are generally disposed in one plane and in
some species the terminal cells of the row differ considerably from
the central cells. There is a wide range of variation in the form of
the cells, which may be ellipsoid, oblong, or fusiform, and the
extremities of all the cells of the coenobium, or of the terminal
cells only, may be furnished with one or more spines. In some the
cells are ridged and in others spines may be
longitudinally
attached to the middle region of the cells. There is a large
parietal chloroplast, often completely filling the cell, and generally
containing a single pyrenoid. Multiplication takes place by the
formation of autospores and autocolonies, and rarely by globular
resting-spores. In cultures these plants have been said to pass
into a palmelloid or gloeocystiform condition.
220 Chlorophycece

Grintzesco 1 has shown that remarkable malformations of

Scenedesmus obliquus can be produced by cultures in a nutritive

medium of agar and glucose, and that this Alga possesses the
physiological property
of liquefying gelatine. He finds a Dacty-
lococcus-st&ge equivalent
to Nageli's D. infusionum, and he
attributes the extensive geographical distribution of this plant to

the ease with which it itself to different

adapts media and different

temperatures.

au.t.

Fig. 92. A, Scenedesmus obliquus (Turp.) Kiitz. from Bradford,

,
W. Yorks. B, the
state of S. obliquus known as D/ictylococcus infusionum Nag., from Bowness,
Westmoreland. C, S. bijiu/atiis (Turp.) Kiitz., from various localities. P, D
,S. <inadriciintlii (Turp.) Breb., from Bradford, W.
Yorks. G, S. quadricauda
var. horridu* Kirchn., from S.E. Surrey. H, S. quadricauda var. maximus
&
W. G. S. West,from Pilmoor, N. Yorks. I K, S. denticulatus Lagerh. var.
linear is Hansg. I and J, from Westmoreland K, from Mayo, Ireland. L, S. spi-
;
;

catus W. & G. S. West, from Saltburn, N. Yorks. (All x 520.) aut, autocolonies.

There are about 10 British species, of which S. quadricauda (Turp.) Breb.

(fig. 92 D F), S. bijugatus (Turp.) Kiitz. (
= 8. obtusus Meyen fig. 92 C) and ;

S. obliquus (Turp.) Kiitz. ( = S. acutus Meyen fig. 92 A and B)

;
are general and
abundant. All three species vary much in size and also in general characters ;

but occur most

they are regular constituents of the freshwater plankton,
in stagnant water, especially in association with Pediastrum
abundantly
Boryanum, Coelastrum
sphcericum, etc. The cells of the largest forms of
S.quadricauda reach a length of 30 /u
and a breadth of 14 p. S. denticulatus

1
Grintzesco, 'Recherch. Experiment, sur la Morph. et la Phys. de
Scenedesmus
Bull, de 1'Herb. Boissier, 2me. ser. 1902.
 Protococcacefp 221

but rarely
Lagerh. var. linearis Hansg. (fig. 92 I K) is widely distributed,
occurs in such abundance as the three preceding species. The cells of S. cos-
tatus Schmidle and S. acutiformis Schroder possess prominent longitudinal

ridges. S. and S. gmnulatus W. & G. S. West are very

Hystrix Lagerh.'
uncommon ornamentation.
species remarkable for their external

Genus Dimorphococcus A. Br., 1849. The cells are arranged

in composite colonies, each colony
consisting of an irregular agglome-
ration of definite groups of four
cells. The of each group are
cells

disposed obliquely in one plane and

are of two kinds; the two central
cells are ellipsoid or oblong arid the
two outer cells are sublimate. The
groups of four are held in position
by irregular portions of the old walls
of the mother-cells. There is a large

parietal chloroplast with one pyre-

noid. Autocolonies of four cells are
produced in each mother-cell these ;

remain attached to the parent-colony

until the latter becomes too large,
when it breaks up into several Fig. 93.Dimorphococcus IIIIKI-
tu$ A. Br. from the plankton
A,
smaller colonies. ;

of Loch Mor Bharabhais, Lewis,

Outer Hebrides; B, from Bowness,
D. lunatus A. Br. is a rare plant in Westmoreland x 520). (

Britain 1 . sometimes occurs in the

It

plankton, but is met with more often in the small tarns of mountainous dis-
tricts. The colonies are 57 86 /z in diameter and the cells 11 25 /* in
length; fig. 93.

Genus Ankistrodesmus Corda, 1838 ; Ralfs, 1848 ; Archer,

1862. [Rhaphidium 1845; Schroderia Lemmermann, 1898.]
Klitz.,
The cells are fusiform with acute apices, rarely obtuse, and they
are generally many times longer than their diameter. They are
straight, lunate, arcuate, or sigmoid, and although frequently
more often variously grouped in loose aggregates.
solitary, are
In some forms the apices are greatly produced and almost bristle-
like. The cell- wall is very thin and there is a single parietal
chloroplast, usually occupying
the greater part of the cell-cavity.

1
Vide West in Journ. Boy. Micr. Soc. 1892, p. 735, t. ix, f. 39; West & G. S.
West in Trans. Hoy. Irish Acad. xxxii, sect. B, part i, 1902, p. 66.
 222 CMorophycece

Pyrenoids are commonly absent, but one or two may occur in

some forms. Multiplication takes place by means of autospores
formed by the division of the contents of the mother-cell they ;

generally arise by oblique divisions, but may be produced by

repeated transverse or longitudinal divisions.
There has never been any doubt concerning the identity of
Ankistrodesmus Corda and Rhaphidium Klitz., but this fact
only seems to have been acted upon by Ralfs in his British
'

Desmids' and Archer 1 when he described his Ankistrodesmus

acutissimus. I am indebted to Dr Nordstedt of Lund for

furnishing me with full information concerning Ankistrodesmus

from Corda's scarce memoir 2
The remarks made by Corda
.

"
under the
heading of Ankistrodesmus novum genus," and
the description and figures he gives of A. fusiformis, are quite
sufficient to establish this genus, and also to show that
Klitzing's
later genus Rhaphidium is identical with it. Kuntze 3 endeavoured
to prove that these two genera should be placed as synonyms of
Micrasterias Corda (1835) 4 a genus which he tried to establish
'

,'

on the assumption that Micrasterias Ag. (1827) should be

relegated to Helierella Bory (1826). Nordstedt 5 has clearly
'
shown that the name Helierella cannot be used as a generic
'

name, and in consequence Micrasterias Ag. (1827) is a valid genus.

It follows from this that Ankistrodesmus Corda is the earliest
generic name given to the common plants which Klitzing named
Rkaphidium, and as Corda's description and figures are also suffi-
ciently characteristic, his name must be accepted for the genus.
The genus Schroderia was established by Lemmermann in
order to include an Alga which had previously been found by
Schroder and described by him as " Reinscltiella? setigera." This
7

Alga I have observed from North Yorkshire along with other

plankton forms. The only distinction that can be drawn between
tichroderia and Ankistrodesmus is the greater attenuation of the

1
Archer in Quart. Journ. Micr. Sci., n. s. vol. 2, 1862, pp. 255 261, t. xii
(in part).
'

Corda in Ahnanach de Carlsbad par J. de Carro, Prague, 1838, pp. 196 198.
2

Kuntze Kevis. Gen. Plant, n, 1891, pp. !)04, 905.

:i

4
Corda in Almanack de Carlsbad, 1835, p. 206. Corda's only mention of
Micrtixtvrids falcnta is in the description of the plates on p. 206; there is no text
reference and the figures are on t. ii, f. 2i).
5
Nordstedt in Hedwigia, 1893, Heft 3, pp. 149151.
B
Leuimermann in Hedwigia, 1898, Bd xxxvii,,p. 311.
7
Schroder, 'Ueber das Plankton der Oder,' Berichte Deutsch. Bot. Gesellsch.
1897, Bd xv.
 Protococcacece 223

apices of the former, and I do not regard this as a sufficient

" "
generic difference. Reinschiella ? setigera must be considered
as an Ankistrodesmus having the same relationship to other
species of the genus as Dactylococcus bicaudatus has to other
species of Dactylococcus.

ou-ts

Fig. 94. A, Ankjj&Mdtisin.Hx falcatu* (Corda) Ealfs, from Bowness, Westmoreland.

B and C, A. falcatus var. acicularis (A. Br.) B, from Pilrnoor, N. Yorks.;
;

C, from the plankton of Loch M. Bharabhais, Lewis, Outer Hebrides. D, A.

falcatus var. tumidus nob., from Puttenhaui Common, Surrey. E, A. falcatus
var. mirabilix nob., from Wimbledon Common, Surrey. F, A. setigerus (Schrb'd.)
nob., from near Bievanlx Abbey, N. Yorks. G and H, A. Pfitzeri (Schrod.) nob.,
from the plankton of Loch M. Bharabhais, Lewis. (All x 520.) uuts, autospores.

As plants of this genus are often very abundant and the synonymy is

much confused, the following synopsis will be found useful :

A. falcatus (Corda) Ralfs. \Micrasteriasfalcata Corda 1835; Ankistro-

desmus fu-siformis Corda 1838 (in part); Rhaphidium fasciculatum Kiitz.
1845 ;
Rh. polymorphism Fresen. var. falcatum Rabenh.] Cells lunate or
arcuate, 16 24 times longer than the diameter (1'5 3 /x), with the apices
acute associated in loose aggregates or close bundles, rarely solitary.
;

Chloroplast visually devoid of a pyrenoid. Fig. 94 A.

Var. acicularis .
\_lihaphidium aciculare A. Br. 1849 Rh. polymorphum
;

var. K<'t'rntre Rabenh.; Closterium subtile Breb. AnJri&trodesmus acutissim.ua

;

Arch. 18(5] Cells usually solitary, commonly straight or slightly curved,

with acutely attenuated apices. Length 36 65 /x breadth 2-5 3'5 /x.
;

Chloroplast sometimes furnished with one pyrenoid (or more). Fig. 94 B

and C. It is most probable that Eh. pyrur/i'iium 'hod. belongs here.
(

Var. duplex \_Rltphi<1itini <////i/e.r Kiitz. 1845.]

. Cells elongate, sig-
moid, associated end to end in pairs, lilt, inrale Chod. may possibly be a
form of this variety.
224 Chlorophycece
Var. tumidus .
[Rh. polymorphism var. tumidum W. & G. S. West
1897.] Cells solitary or in small aggregates, curved, in the middle inflated;

apices most acute; length 61 73 p breadth 4'5 6'5 p. Chloroplast with ;

one or two pyrenoids, or destitute of them. Fig. 94 D.

Var. mirabilis .
[Rh. polymorphum var. mirabile W. & G. S. West
1897.] Cells solitary and considerably longer than in typical A. falcatus,

variously curved, often sigmoid apices very acute. Chloroplast completely;

interrupted in the middle of the cell and often fragmented ; vacuoles in the
cell- protoplasm often containing a single moving corpuscle. Length up to
117 fj. ;
breadth 2 3'5 M .
Fig. 94 E.
Var. spiralis Turn. 1893; Rh. fasciculatum var. spirale
.
\_Rh. spirale
Chod. 1902.] grouped in bundles of 4 or 8, twisted round each other in
Cells
the central region of the cells, but free at their extremities.
Var. spirilliformis .
[Rh. polymorphum var. spirale W. & G. S. West
1898.] Cells always solitary, spirally twisted, making 1 1-J- turns; apices
very acute. Often occurs in prodigious quantity in stagnant water.
The above varieties of A. falcatus are widely distributed and some of them
are abundant. They occur most abundantly in small ponds, but are also
frequent in the freshwater plankton.
A. convolutus \_Rliaphidium convolution Rabenh. 1868.]
-
. Cells
only 3 6 times longer than the diameter, lunate or arcuate ;
.solitary, short,

apices very acute diam. 3 6 p.;

A. Pfitzeri [Rhaphidium Pfitzeri Schroder.] Cells straight, some-

.

what asymmetrical, 5^ 6 times longer than the diameter, much attenuated

with rounded apices forming small colonies embedded in mucilage. Length
;

38 49 breadth 7 8 p.. In the Scottish plankton not uncommon. Fig.

/z ; ;

94 G and H. Perhaps this species would be better placed as a form of A. biplex

(Reinsch) nob. [ = Rh. bipkx Reinsch 1867.]
A. setigerus [Reinschiella ? setigera Schroder 1897; Schroderia
-
.

setigera Lemm. 1898 Rhaphidium sctigerum W. & G. S. West 1901.] Cells

;

fusiform, with the apices much attenuated into tine hair-like prolongations.
Chloroplast with one pyrenoid. Diam. 5 7 6'5 p; length 75 88 p.. Mostly -

found in the plankton. Fig. 94 F.

Genus Closteriopsis Lemm., 1898 This genus only differs 1

.

from Ankistrodesmus in the great length of the cells and in the

consequent increase in the number of pyrenoids. The cells are
narrow, greatly elongated, and the extremities are much attenuated
into bristle-like points. The Chloroplast contains a dozen or more
pyrenoids in an axial series.

Cl. longissima Lemm. is only known from the plankton. In the British
Islands it has been found in
Lough Neagh, Ireland, from Finstown, Orkneys,
and in Loch Asta, Shetlands. Length 330 p. ;
breadth 3'8 4-2 p.

Genus Actinastrum Lagerh., 1882. The cells are ovoid,

oblong or club-shaped, from three to six times longer than their
1
Leminermauu in Forschungsberichten Biol. Stat. Plon. vii, 1898, p. 29.
 Protococcacece 225

diameter, and are generally attached by their apices to form

radiating colonies of small size. Each
cell possesses a parietal

chloroplast furnished with a pyrenoid. Multiplication takes place

by the longitudinal division of the cell-contents, with the occasional
formation of another transverse wall. The products of division
diverge outwards, but remain attached by their proximal ends,
and the colonies sometimes reach a relatively large size owing to
the repeated new divisions of the contents of the radiating cells.
As a general rule four young cells (autospores) are produced in
each mother-cell.
A. Hantzschii Lagerh. is a very rare Alga in Britain and is confined
principally to the plankton. Length ofcells 10 24 p, breadth 3 6 /x.
;

Genus Selenastrum Reinsch, 1867. In this genus the cells

are arcuate or lunate, attenuated to

fine points, and arranged back to back
to form a 4- or 8-celled colony of small
size. The cell-wall is thin and firm,
and the chloroplast is without a pyre-
noid. The multiplication is by auto-
spores which arise as in Ankistrodesmus
and Kirchneriella.
The species of this genus are very un-
common, usually occurring amongst other
water-plants at the margins of ponds and
lakes. S. Bibraianum Reinsch, S. gracile
? Fig. 95. A D, Selenastrum
Reinsch (fig. 95 A
D) and S. acuminatum gracile Eeinsch; A C, from
(fig. 95 E
Lagerh. G) all occur in Britain. near Settle, W. Yorks.; D,
Chodat considers the latter species to be a from Puttenham Common,
7 .,, Surrey. E G, S. acumina-
Scenedesmus, but that I cannot agree with. tum Lagerh from B owness,
.
t

Westmoreland. (All x 520.)

Genus Kirchneriella Schmidle,
1893. [Indus. Selenoderma Bohlin, 1897.] The cells are arcuate
or crescent-shaped, attenuated or subcylindrical, often bent until
the apices almost touch each other they are loosely aggregated,
;

without any definite disposition, within a large enveloping mass of

jelly. The cell- wall is very thin and the chloroplast is parietal,
being situated on the convex wall of the cell. There is one
pyrenoid, but it is frequently absent. The multiplication is by
autospores, four or eight of which are produced in a mother-cell
by oblique or more or less transverse divisions. The genus differs
from Selenastrum in the loosely aggregated colonies of cells, which
W. A. 15
226 Chlorophycece

are irregularly disposed, and in the large enveloping mass of

B
Fig. 96. Kirchneriella obcsa (West) Schmidle. A, B, D and E, from Bowness,
Westmoreland ; C, from the plankton of Loch Mor Bharabhais, Lewis, Outer
Hebrides ( x 485).

K. obesa (West) Schmidle (breadth of cells 2 9'5 p. apices 1-5 4 /* apart;

;

greatest diameter of cell 6 16 p.; fig. 96 A E) and K. lunaris (Kirchn.)

Mob. are widely distributed in the British Islands, occurring sparingly in the
small ponds and lakes, but in quantity in the plankton of the larger lakes.

Sub-family IV. OOCYSTJDE.E.

This sub-family is characterized by the globose or ellipsoid cells
(curved or even sublunate in Nephrocytivm), which are frequently
retained within the swollen wall of the old mother-cell. There
may be one or many chloroplasts in each cell, which are parietal
and usually contain one pyrenoid. In all except Palmellococcus
the cell-walls are firm and somewhat thick. The multiplication is
by autospores, which generally grow and attain their full size
whilst still forming part of the mother-colony.
The following are the British genera :

* Cells Oocystis.
ellipsoid
** Cells curved, subcylindrical or sublunate Nephrocytlnm.
*** Cells
spherical.
t Cells large, solitary and free-floating Eremosplmra.
tt Cells minute, forming a thin stratum Palmellococcus.
t tt Cells minute, sparsely scattered and symbiotic Chlorella.
 Protococcacece 227

Genus Oocystis Nag., 1845. The cells are ellipsoid, sub-

cylindrical or panduriform, with a firm cell-wall which commonly
possesses a nodular thickening at each pole. There are usually
several parietal chloroplasts in each cell, destitute of pyrenoids.
In some cases, however, there may be one pyrenoid in each
chloroplast. Multiplication is
by autospores, which are generally
retained for some time within the greatly swollen wall of the
mother-cell. Sometimes several generations are contained within
one large mother-cell-wall.

Fig. 97. A and B, Ooci/stis solitaria Wittr., from Ben Lawers, Perthshire. C and
D, O. cmssa Wittr. C, from Lanlivery Moor, Cornwall D, plankton form
; ;

from Lough Beg, Ireland. E and F, 0. panduriformis W. & G. S. West; E,

from Pilmoor, N. Yorks.; F, from near Clifden, Ireland. G, 0. elliptica West,
from Derryclare Lough, Galway, Ireland. (All x485.)

There are about ten British species of the genus, some of which are widely
distributed in the quiet waters of ponds and lakes. 0. solitaria Wittr. (length
of cells 15 48 ^ breadth 9'5 25 /^; fig. 97
;
A
and B) is undoubtedly the
commonest species, although 0. elliptica West (fig. 97 G) is abundant.
0. parva West & G. S. West
(length 6 12 p breadth
;
/*)
47
is the smallest

species and 0.giyas Arch, (length 41 50'5 breadth 32'5 40 p.) is the
/j. ;

largest. 0. crassa Wittr. is not common except in the plankton fig. 97 C

;

and D. 0. asymmetrica W. & G. S. West is another solitary species.

Chodat states that 0. giyas Arch, and 0. panduriformis W. & G. S. West
(fig. 97 E and F) are merely stages in the life-history of JSremosphcera, but
this statement I do not accept. These plants often occur in localities from
which EremosphcBra is absent and they reproduce themselves in the manner
of true species of Oocystis. Moreover, I invariably find the life-history of
Eremosphcera to be remarkably free from polymorphic forms (vide page 229).
152
228 Chlorophycece
Genus Nephrocytium Nag., 1849. The cells are oblong,
ellipsoid or subcylindrical, slightly curved or sublimate, sometimes
almost reniform. There is at first a large expanded chloroplast in
each cell, furnished with a single pyrenoid, but later the chloroplast
fragments. The multiplication is by autospores similar to those
of Oocystis, and formed within the mother-cell soon after the

segmentation of the chloroplast. The young autospores are often

spirally disposed round the inside of the mother-cell-wall. The
genus is distinguished from Oocystis primarily by its curved cells
without any trace of apical thickenings.

wo
Fig. 98. A, Nephrocytium obesum West, from Angle Tarn, Cumberland. B, 2V.

ecdyxiscepanum West & G. S. West, from near Goring, Oxfordshire. C F,

N. hinatum West; C E, from near Bowness, Westmoreland; F, from near
Roundstone, Galway, Ireland. (All x 367.)

N. Agardhianum Nag. (inclus. N. Nagelii Grun.) is a widely distributed

species in the stagnant waters of small pools and lakes; length of cells 12
22 p.; breadth 7 12 p. N. obesum West is the largest species, characterized
by the short, stout and by the great thickness of the mother-cell-walls
cells ;

length of cells 34 42 /x breadth 24 28 ^ fig. 98 A. N. lunatum West is a

; ;

characteristic species (supposed by Chodat to be a stage of T. Agardhianum)

which is very local, but sometimes abundant among submerged Sphagnum ;

length of cells 14 18 /* breadth 4 6 '5 p tig. 98 C F.

; ;
N. ecdysiscepanum
 Protococcacece 229

W. & G. S. West is a curious species in which the mother-cell-walls throw off

a number of integuments, several generations being disposed in a fan-shaped
manner and held in position by the partly exuviated layers of the mother-cell-
walls; length of cells 24 26'5 p. breadth 1317 /* fig. 98 B.
;
;

Genus Eremosphsera De Bary, 1858. [Chlorosphcera Hen-

trey, 1859.]
The cells are solitary, large, and spherical, with a
thick, firm cell-wall differentiated into two layers. Each
distinctly
cellcontains a large number of small parietal chloroplasts furnished
with a conical projection directed towards the centre of the cell.
There is one in each chloroplast.
pyrenoid
The nucleus is gene-
rally contained
in a small mass of protoplasm suspended in the

Fig. 99. Ervmuxplucru viridis De Bary, from near Clapham, W. Yorks. (

x 175).

central region of the cell by a network of protoplasmic threads.

of a
Multiplication takes place by the division of the contents
mother-cell into two or four smaller but similar daughter-cells
(autospores), which are set free by the rupture of the mother-cell-
wall. Chodat 1 has described the occurrence of certain polymorphic
forms of this Alga, but although I have examined large quantities
of it from every part of the British Islands, and from elsewhere,
I have never yet seen any trace of such forms. Specimens kept
under cultivation for two years developed no forms other than
globular daughter-cells. Moore 2 also disputes the alleged poly-
morphism of this Alga.
E. viridis De Bary is widely distributed all over the British Islands, more
especially in Sphagnum-bogs. It is a constant associate of certain Desmidx.
The cells vary from 55 200 p. in diameter; fig. 99.
Genus Palmellococcus Chodat, 1894. \JProtococcus Ag. 1824
(in part).] The cells are more or less globular, with a firm cell-
wall, and aggregated to form a thin mucous stratum.
There is in
1
Chodat in Botan. Zeitung, liii, 1895, t. v.
2
Moore in Proc. Amer. Assoc. Adv. Sci. 1900, pp. 278, 279.
230 ChlorophyeecB
each cell a parietal plate-like chloroplast devoid of a pyrenoid,
which is often hidden by an orange-
red oil. There are three methods of
multiplication 1st, by division of the
;

original mother-cell into two or four

daughter-cells ; 2nd, by a rejuvene-
scence of the cell-contents and an
exuviation of the wall of the mother-
p lm * l oc ccusmini -
-
[
atus (Kutz.) Chodat,
.

trom near by the formation of 8,

cell; 3rd,
Bradford, W. Yorks. (x720). 64 spores within the wall
16, 32, or
of the mother-cell, which then rup-
tures and sets them free. Palmellococcus differs from Pleurococcus
in the absence of pyrenoids from the chloroplasts, in the methods
of reproduction, and therefore in the different aspect of the cell-

aggregates.
(Pleurococcus miniatus Nag.) is not an in-
1
F. miniatus (Kiitz.) Chod.
frequent Alga on the outer surfaces of plant-pots and similar objects, forming
a moist, brownish-green scum, which often turns to an orange-red colour.
Diam. cells 212-5 p. ; fig. 100.
Certain plants belonging to this genus have in the past been referred to
"
Protococcus Ag." The latter genus is obsolete, having included Algte which
are now referred to a number of other genera.

Genus Chlorella Beyerinck, 1890 2 . The cells are small,

globular or ellipsoid, with firm cell-walls, and with a single parietal

chloroplast containing a pyrenoid. They occur in abundance in
symbiotic relationship with Hydra viridis, species of Amoeba,

Paramcecium, Ophrydium, etc. Multiplication takes place by

the quadri partition of the cell contents. Radais 3 has confirmed
Beyerinck's observations that this Alga has the faculty of cell-
increase and the formation of chlorophyll in the dark as in the

light. Grintzesco 4 also affirms that development takes place more

rapidly in total darkness than in full daylight. The latter author
has cultivated Chlorella in various media and finds that glucose
stimulates its development and that peptone is a better source
of nitrogen than nitrates. Cultures of this Alga do not liquefy
gelatine.
Clil. r/il(/(tris Beyr. is widely distributed and often abundant in cultures
or in stagnant aquaria. The cells are 5 10 in diameter.
fj.

Chodat in Bull. Herb. 1894, pp. 429 and 599.

1
Boissier, torn, ii,
2
Beyerinck in Botan. Zeitung, xlviii, 1890.
3
Badais in Comptes Bendus, cxxx, 1900, p. 793.
4
J. Grintzesco in Bev. Gen. Bot. xv, 1903.
 Protocoecacece 231

Sub-family V. TETRAEDRIE^.
The plants of this sub-family are always solitary unicells.
Each cell is flattened and angular, usually with a definite number
of angles, or it is tetrahedral, octahedral, or polyhedral. The
angles may be rounded, emarginate, or furnished with spines.
Genus Tetraedron Kiitz., 1845. [Polyedrium Nag., 1849.]
The cells of this genus, which occur as solitary individuals or
rarely collected into temporary aggregates, are flattened and
angular (triangular, quadrangular,, or polygonal) or polyhedric.
The angles are obtuse, acuminate, or furnished with one or more
simple or furcate spines. There is a single large chloroplast,
parietally disposed and containing one pyrenoid. Multiplication
takes place by the formation of four or eight autospores, which are
set free by the rupture of the wall of the mother-cell. Sometimes
they are expelled into a delicate vesicle, which, however, soon
disappears.

Fig. 101. A, Tetrac.dron minimum (A. Br.) Hansg., from Keston Common, Kent.
B, T. caudatum (Corda) Hausg., from Pilmoor, N. Yorks. C, T. reyulare
from near Bowness, Westmoreland. I),
Kiitz., T. enorme (Ealfs) Hansg., from
Mickle Fell, N. Yorks. E G, T. Iwrrlditni W. & G. S. West, from Putney
Heath, Surrey. (All x 450.) ants, autospores.

There are about 14 species of the genus known to occur in the British
Islands. T. minimum (A. Br.) Hansg. is the most abundant of the flattened

species; diam. of cells 6'5 16 /^;

thickness of cells 5 7 p.; fig. 101 A.
T. regulare Kiitz.(
= Polyedrium
tetraedricum Nag.) is the commonest of the

polyhedric species; diam. of cells 13'5 40 p. fig. 101 C. ;

T. enorme (Ealfs)
Hansg. was originally described as a Desrnid tig. 101 D. ;

The genus Cerasterias Reinsch (1867) is sometimes separated

from Tetraedron owing to the depth of the lobulation. The cells
are solitary, tetrahedric in character, and the divisions into lobes
232 Chlorophycece

are so deep that there is no central body. C. rhaphidioides Reinsch

and C. longispina (Perty) W. & G. S. West are both plants of

rare occurrence.

Sub-family VI. PHYTHELIE^.

This one of the most interesting sub-families of the Pro-
is
in it have only recently been
tococcacese, and the Alga? contained
the of
brought to light, largely by plankton investigations
Lemmermann and Chodat. The plants are unicellular or some-
times grouped so as to form a more or less definite coenobium,
and in all cases they float freely in the water. As a rule the cells
are almost devoid of a mucous envelope, and they are furnished
with several stiff considerably longer than their
bristles own
occurs the formation of
diameter. Multiplication typically by
which attain all the characters of the adult
autospores, usually
before their liberation from the swollen wall of the mother-cell.

Zoogonidia have been observed

in Golenkinia.
The name of the sub-family is derived from the genus Phythelios
Frenzel (1891), an Alga which was originally described as a
Heliozoan. All the genera are practically confined to the plankton
of large lakes, although a few of them are occasionally observed in
the surface waters of ponds. The long bristles of these Algae are
as a result of a free-floating
protective characters developed
existence amidst numerous animals to which they would other-
wise be an easy prey.
The genera have been well worked out and monographed by
Lemmermann 1
. Four of them are known from Britain.
A. Cells globular.
* Cells solitary with evenly distributed bristles Golenkinia,
** Cells in colonies of 8, 16, or more, bristles attached
to the outer faces only Richteriella.

B. Cells ellipsoid or subcylindrical.

* Bristles with a basal swelling Lagerheimia.
** Bristles without a basal swelling Chodatella.

Genus Golenkinia Chodat, 1894. The cells are globular,

which is enveloped in a
usually solitary, with a firm cellulose wall,
thin layer of mucilage. Each cell is furnished with a number of
over the
radiating bristles of considerable length, evenly disposed

1
Lemmermann in Hedwigia, Bd xxxvii, 1898.
 Protococcacece 233

outer surface of the cell. There is a parietal chloroplast con-

taining one pyrenoid. Multiplication normally takes place by

autospores, but also by simple vegetative division, and reproduction
by quadriciliate zoogonidia has been observed by Chodat.
G. radiata Chodat 1 is known from Surrey ;
diarn. of cells 10 15 //. ; length
of bristles 2545 /z ; fig. 102 I) and E. G. paucispinosa West & G. S. West 2
is known from the plankton of Lough Neagh, Ireland ;
diam. of cells 15 16 p. ;

length of bristles 16 /*; fig. 102 F.

Fig. 102. A, Ricliti'i-iella botryoidcs (Schm.) Lemm., after Lemmermann, x 520.

B and forma quail risetti (Lemm.) Chod. B, from the plankton
C, R. botryoides ;

of Lough Beg, Ireland (x450); C, after Lemmermann x 520). D and E, (

Golenkinia radiata Chod., after Chodat ( x about 800). F, Golenkinia pauci-

W. & G. S. West, from the plankton of Lough Neagh, Ireland ( x 450).

Genus Richteriella Lemmermann, 1896. The cells are spherical

and generally aggregated to form loose ccenobia of 8, 16, 32, or 64

cells. The cell-wall is thin and firm, and is furnished with long,
radiating bristles attached only to those surfaces of the cells which
face outwards. a single parietal chloroplast with one
There is

pyrenoid. Multiplication has only been observed to take place by

vegetative division. This genus only differs from Golenkinia in
the aggregation of the cells and the fact that the bristles are more
or less confined to the exposed surface of each cell.

1
Chodat in Morot, Journ. de Bot., Paris, 1894, p. 305, t. iii.
2
West & G. S. West in Trans. Koy. Irish Acad. xxxii, sect. B, part i, 1902,
p. 68, t. i, f. 18.
234 Chlorophycece
R. botryoides (Schmidle) Lemm., forma quadriseta (Lemm.) Chod. is
known from the plankton of Lough Beg, Londonderry, Ireland; diam. of
cells 39-6 p. ; length of bristles 2360 p ; fig. 102 B and C.

Genus Lagerheimia Chodat, 1895. The cells are solitary,

ellipsoid, or
subcylindrical with
rounded extremities, and with a
firm cell-wall. There are four bristles

arranged in diverging pairs at each

pole, or disposed
one at each pole
and two equatorially.
Each bristle

has a wart-like thickening at its

base. There is
single parietal a

chloroplast with one pyrenoid.

The
is by autospores.
multiplication
L. subglobosa is known from
Lemm.
Lough Gartan, Donegal, Ireland (diam. of
cells 5-59-4^; length of bristles 10'5
13 p.; fig.
103 D and E), and L. gen&vensis
Chod. from the south of England (diam. of
cells 3 p. ; length of cells 910 p. ; length of
bristles up to 16 p. fig. 103
;
A C).

Genus Chodatella Lemmermann,

1898. This genus only differs from
Fig. 103. A C, Lagerheimia the preceding one in the absence of
genevenxis Chod., after Chodat ( x the swellings or wart-like thicken-
about 850). D
and E, L. sulic/lo-
bosa Lemm. D, after Lemmermann
;
ings at the base of the
bristles.
(x520); E, from Lough Ga.rtan, The cells are solitary, ellipsoid, and
Donegal, Ireland x 450). F and (

G, Chodatella breviseta W. & G. S. furnished with four or many elon-

West, from Lough Gartan, Ireland
H and I, Ch. ciliata gated bristles, which are sometimes
(x450).
(Lagerh.) Lemm. var. amphitricha radiating sometimes curved.
and
(Lagerh.) Chod. ( x 450) H, from
There may be one or several parietal
;

Skipwith Common, E. Yorks. I, ;

from near Bowness, Westmoreland. or without pyre-

chloroplasts, with
auts, autospores.
noids. The genus differs from Oocys-
tis in the absence of the polar thickenings and the presence of the
long spine-like bristles.
Ch. clliata (Lagerh.) Lemm. var. amphitricha (Lagerh.) Chod. [
= Ch.
radians (West) Lemm.] occurs in several localities in the British Islands;
of bristles 12 20 p.; fig.
length of cells 8 18 M breadth 4 13'5 p.; length
;

103 H and I. Ch. breviseta West & G. S. West is known from Lough Gartan,
breadth 8 9-5 /*; length of
Donegal, Ireland; length of cells 12 12-5/*;
bristles 11 '5 17 '5 p.; fig. 103 F and G.
 Protococcacece 235

Sub-family VII. DICTYOSPILERIE^E.

This sub-family contains a few genera the affinities of which
are somewhat doubtful. The cells are globose, ovoid, or ellipsoid,
and are associated to form more or less indefinite colonies. The
colony is often of a fragile nature, the cells being held in position
by the persistent old walls of the mother-cells, which sometimes
become transformed into definite connecting-threads. A copious
mucous investment is present in some, but in others it may be
entirely absent. The multiplication is by simple vegetative divi-
sion or by 'the formation of four daughter-cells (autospores) within
the wall of the mother-cell, which gradually splits open and permits
their escape. Reproduction by biciliated zoogonidia has been
observed by Zopf and by Massee 1
in Dictyosphwrium.
The sub-family is most probably an
artificial one and perhaps
it should not have a place in the Protococcacege.

The five following genera are British :

A. Cells indefinitely disposed.

* With
well-marked, subdichotomous connecting-
th reads ;
chloroplast parietal Dictyosphcerium.
** Cells in threads
radiating series; connecting
scarcely visible; chloroplast axile Dictyocystis.
B. Cells grouped in fours in one plane ;
colonies ir-

regular Tetracoccus.
C. Cells in botryoidal clusters.
*
Freely exposed in a thin gelatinous envelope Botryococcus.
** Clusters covered by a firm, irregular, tough
membrane Ineffigiata.

Genus Dictyosphserium Nag., 1849. The cells are globose,

ovoid, or subreniform in shape, with a firm cell-wall, and they are

connected by dichotomously branched
^3 <P=SH rv\ threads to form a roughly spherical
or ellipsoidal colony. The entire
colony is enveloped in mucus, and
the cells are situated somewhat far
apart towards its periphery, large
colonies often becoming very irregu-
Fig. 104.
Dictyosphteriumpul-
chelium Wood. A, from the plank- lar. Each cell contains a more or
ton of Loch Shin, Sutherland; B, i n j i i i . i i

from Cam Fell, W. Yorks. x 450. less bell-shaped, parietal chloroplast,

1
G. Massee in Journ. Linn. Soc. Bot. xxvii, 1891.
236 Chlorophycece

furnished with a single pyrenoid. The connecting-threads are

often derived from the old mother-cell-walls, but in some cases it
is doubtful if do
theyarise in this manner. Multiplication takes
place normally by the
formation of four daughter-cells within the
but
mother-cell. Reproduction by biciliated zoogonidia occurs,
has been very rarely observed.

D. Ehrenbergianum Nag. is a widely distributed British Alga, often

occurring in quantity in the surface waters of ponds
and in the plankton of
lakes diam. of cells
;
410
/*.
D. pulchellum Wood (fig. 104), D. reniforme
Buln. and D. oviforme Lagerh. are more rarely found, but it is probable that
the five so-called species of this genus are merely forms of D. Ehrenbergianum.

Genus Dictyocystis Lagerh., 1890 The cells are ellipsoid

1
.

or oblong, and are arranged in radiating series to form a small

free-floating colony. The radiating series of cells frequently

branch and the cells are held in place by delicate mucous threads.
Each cell possesses a central chloroplast with one pyrenoid.
D. Hitchcockii (Wolle) Lagerh. is a rare British Alga, occurring in the bogs
of N. Ireland and N. W. Scotland, and also in the Scottish plankton. Diam.
of cells 9 11 /x; the American specimens are larger.

Genus Tetracoccus West, 1892 s [Westella De Wild., 1897

.

(in part).] The cells are small, globose or subglobose, sometimes

a little angular, and are closely
arranged in groups of four. These
groups are connected by the old
mother-cell-walls into free-floating
colonies of small size, consisting
of a maximum number of about
80 cells. The four cells of each
C
group are disposed in one plane,
Fig 105 Tetracoccns botry aides
d the o]d wallg of the mother-
West, from Bowness, Westmoreland.
A and B, x450; B and C, two "te- cells become transformed into deli-
trads," x7l5. cat e There
connecting-threads.
is one chloroplast which contains several large granules, but the
demonstrated.
presence of pyrenoids has not yet been definitely
takes place by the formation of four daughter-cells
Multiplication
within the mother-cell, which arise by the division of the cell-
contents in two directions in one plane. The colonies are almost
entirely free from enveloping mucus.

Lagerheim in Nuovo Notarisia, 1890, p. 226.

1

2
West in Journ. Boy. Micr. Soc. 1892, p. 735, t. x, f. 4348.
 Pi -o tococcacecp 237
'

Schmidle gave what he termed an amended description of

'
1

this genus in 1894, but as the plant he included in it is not a

amended is not a correct
species of Tetracoccus, his description
one 3 .

T. botn/oides West is widely distributed in the British Islands, generally

of large lakes.
occurring in the surface waters of ponds and in the plankton
Diam. of cells 3-8 57 /* diam. of colonies 3057 p; fig. 105.
;

Genus Botryococcus Klitz., 1849. The colony is free-floating

and an aggregate of botryoidal groups of
consists of cells. The
cells are globose or ovoid in form
and are closely aggregated to form
clusters of 16 or 32 cells, the clusters

being held together partly by old

mother-cell-walls and partly by a

gelatinous investment. There is a

single cup-shaped chloroplast in each

cell, pyrenoids have not been
but
106.
. , Fig. Botryococcus
observed. (Jnodat and Uretier 8 nave Bmunii Kiitz., from the New
observed in the chloroplast a small Hants A small colony;
forest,
B, two isolated cells (
x 450).
body which can be regarded as a
pyrenoid without an amylosphere. As a rule this Alga is of a
bright green colour, but when occurring in large quantity, as it
frequently does in the freshwater plankton, the cells become filled
with a brick-red oil.

B. Brauitii Kiitz. is the best known representative of the genus, and is a

frequent plant in bog-pools, large ponds, lakes, etc. Diam. of cells 5'5 9 p ;

fig. 106. B. gtideticus Lemm. (which may only be a form of B. Bran nil with
globose cells) and B. calcareus West are rarer British species.

Genus Ineffigiata West & G. S. West, 1897; em. 1903.

This Alga consists of free-floating colonies of very irregular form
and destitute of a gelatinous investment. The colony is composed
of several families of cells agglutinated together, each family

being small, more or less spherical,

and consisting of a peripheral
layer of cells surrounding
a central cavity. The cells are ellipsoid
or ovoid in form, and each one is furnished with a parietal ehl<>n>-

plast, often containing

what has been described as a single small

1
Schmidle in Flora, 1894, Heft 1, p. 45.
2
Vide West & G. S. West in Journ. Roy. Micr. Soc. 1896, p. 162.
3
Choclat & Cretier in Arch. Sci. Phys. et Nat. x, 1900.
238 Chlorophycew

pyrenoid, but which I am inclined to believe is a small granule of

starch. In some cells starch appears to be quite absent. The
outer surface of each family of cells is enveloped in a tough elastic
membrane of irregular form,
which contains a trace of
cellulose, and is folded and
produced into all manner of

irregular wrinkles, lobes, pro-

cesses and spines. Sometimes
these irregular projections are
wanting, but at other times
they are exceedingly nume-
rous. The membrane is a
secretion of the underlying
cells, and its presence renders
observations on this Alga
more difficult than on any
other of the Protococcacese.
Sometimes the smaller colo-
nies are united by more or

prolongations of the
less rigid

enveloping membranes into

much larger colonies.
The families multiply by
107. Ineffigiata neglecta W. & division, forming larger and
G. S. West, from Harris, Outer Hebrides.
larger colonies, which ulti-
A, outline of colony B, smaller colony
; ;

C, part of single family in section; D, mately become separated into

showing escape of cells from a small smaller groups by the develop-
colony.
ment of elongated processes
of the enveloping membranes. The reproduction is unknown 1 .

In situations in which Ineffigiata occurs in quantity, such as

in the freshwater plankton, the cells develop the brick-red oily
material which is found in the
preceding genus.
/. ncglecta W. & G. S. West is one of the most widely distributed of British
Algie, occurring in all kinds of situations in ditches, bogs, tanks, water-

butts, etc.and forming a regular and considerable constituent of the fresh-

water plankton. Diam. of single families 21 56 ^, of colonies 46 350 p ;

length of cells 5'7 10'5 M breadth

,
3'4 5'3 /* ; fig. 107.

1
Vide Journ. Bot. March, 1903, t. 447, f. 16.
 Pftlmdhtcece 239

Family 8. PALMELLACE^.
The Palmellaceaeone of the most primitive families of green
is

Alga?, primarily distinguished from the other groups of the

Protococcoidea? by the indefinite colonies of cells enveloped in a

conspicuous mass of mucilage. In the ordinary vegetative con-

dition these plants present the appearance of a group of more
or less irregularly disposed cells embedded in a copious mass of

jelly, which is either structureless or differentiated into concentric

envelopes. The colonies are either microscopic or macroscopic,

and sometimes reach a length of several centimetres.
The cells are globose or ellipsoid, of small size, and are

frequently arranged in pairs or in groups of four. Sometimes

' '

these groups of four or tetrads are disposed in a tetrahedral

manner, but at other times they are situated in one plane. Each
cell contains a somewhat bell-shaped chloroplast which may or

may not be furnished with a pyrenoid. The nucleus is situated

in the hollow of the chloroplast. In some genera (e.g. Tetraspora
'

and Apiocystis} each cell is furnished with a pair of pseudocilia,'

which consist of long motionless protoplasmic threads penetrating
through the enveloping mucus to the exterior. These were first
discovered by Thuret.
Multiplication takes place by cell-division in two or three
directions, followed sooner or later by a diffluence of a large part
of the enveloping mucus and the consequent dismemberment of
the colony into smaller portions, each of which increases as before
either by simple cell-fission or by the formation of two or four

daughter-cells within each mother-cell.

Asexual reproduction takes place by biciliated zoogonidia.
The formed either by the transformation of a vegetative
latter are
cell into a zoogonidangium in which several zoogonidia arise, or

by the assumption by the ordinary vegetative cell of the motile

Chlamydomonadine condition. The motile state greatly resembles
the Chlamydomonad-type, and these plants have no doubt arisen
by the intercalation of a simple though well-marked vegetative
condition between two successive motile phases.
Sexual reproduction has been observed in some of the Palmel-
lacese. It consists of a fusion of isogamous planogametes, either
similar in all respects to the zoogonidia and produced singly in a
 240 Cliloropliycem

gametangium, or much smaller than the zoogonidia and produced

in numbers from a gametangium.
In some of these plants the colony has no definite form, but in
others the cells, although irregularly grouped, are contained in a
mucous mass which invariably assumes a definite shape.

The family can be divided into three sub-families, in each of which the
mucus may be indefinite or developed in accordance with some definite plan.

Sub-family I. Palmellece. Cells irregularly grouped within a

structureless mass of mucus.
Sub-family II. Tetrasporece. Cells grouped in fours or sometimes
irregularly disposed at the periphery of a structureless mass of mucus.
Each cell with two pseudocilia.

Sub-family III. Glceocystidece.. Cells grouped in twos or fours

within a lamellose mucous investment.

Sub-family I. PALMELLECE.
This sub-family
is characterised
by the large number of globose
cellswhich are irregularly grouped within a structureless mass of
mucus. The latter is usually of indefinite extent, but in Palmo-
dactylon it is more or less cylindrical and often much branched.
The cell-walls are generally firm and thin, and in Schizochlamys the
outer layers are periodically thrown off in one or several pieces.

Genus Palmella Lyngb., 1819. The cells are spherical, with

thin cell-walls, and they are surrounded by mucous coats which
have fused to form an indefinite mass of jelly. The parietal
chloroplastcontains a pyrenoid. Multiplication takes place by
repeated bipartitions of the cells in all directions of space, ac-

companied by an extreme gelatinization of the mother-cell-walls.

Reproduction occurs by macrozoogonidia, by microzoogonidia, and
by small isogamous planogametes.
One of the few true species of this genus is P. miniata Leibl., which occurs
as amucous expansion of a brick-red colour on damp ground, wet rocks, etc. ;

diarn. of cells 3 P. mucosa Kiitz. and P. hyalina Breb. are aquatic

5 p.

species of a green colour the former may possibly be a state of Tetraspora.

;

Genus Palmodactylon Nag., 1849. This is a well-marked

genus of the Palmellea?, differing from Palmella mainly in the
definite form of the enveloping jelly. The chloroplast is parietal and

irregularly lobed, but is destitute of a pyrenoid. Multiplication

of the cells takes place in all directions, but preponderates in
 Palmettacece 241

one direction, so that the cells are grouped irregularly within

a cylindrical mass of mucus. This cylindrical mucous envelope
frequently branches, each
branch being similar to the
_ocpa, ,M,n2#><?v
primary mucous cylinder.
P. rarium Nag. (inclus. P.
subramosum Nag.) is a common
British Alga in peaty ditches
and in bog-pools. Diam. of cells
6'5 9'5 ju.; diam. of cylindr.
mucous investment 11 "5 33 /x ;

fig. 108.

Genus Schizochlamys
A. Br., 1849. The cells are
globular or slightly ellipsoid,
with a firm cell-wall of some
thickness, and they are ir-

regularly disposed within an

indefinite j elly. The chloro-
plast contains no pyrenoid.
The outer portion of the Fig. 108. Palmodactylon vnriuni Nag.,
A C, from Esber Common, Surrey; D,
cell-wall commonly becomes from Strensall Common, N. Yorks. A, x 100;
ruptured, being cast off
B D, x450.
either in one piece or in
four distinct pieces, which
are held in a position some-
what remote from the cell

by the intervening jelly.

It is the formation of this \J
large quantity of gelatinous
material that causes the
Fig. 109. A B, Scliizochlamys (jelatinosa
rupture of the firmer part A. Br. from Loughrigg, Westmoreland. C,
,

of the cell-wall. -SV/f. (Idicatula West, from near Bowness,

Multipli-
Westmoreland x 450).
cation occurs by division (

into two or four daughter-cells,

usually subsequent to the rupture
of the outer layer of the cell-wall.

>'.
i/i-lutinosa A. Br., in which the outer layer splits in four pieces, occurs
in ditches, ponds, etc. ;
diam. of cells 9'5 15 /x; fig. 109 and B. S. deli- A
catula West is also a frequent species, in which the outer
layer is thrown off
in one piece ;
diam. of cells 5'8 6'7 /x ; fig. 109 C. S. yelatinosa usually

w. A. 16
242 CMorophycece
occurs in very extensive gelatinous masses, often several cms. in diameter,
whereas the colonies of S. delicatula are very much smaller, rarely exceeding
a diameter of 300 /^. The latter species has also a distinct preference for
Sphagnum-pools.
Genus Sphaerocystis Chodat, 1897 l
. The
are globose cells
and sparsely aggregated within a free-floating globular mass of
very transparent jelly. The number of cells within each gelatinous
sphere varies from 1 to 16 (or more), and they are usually disposed
towards the periphery. Sometimes there is a slight indication of
lamellation in the jelly immediately surrounding the cells. The

-
.

Fig. 110. Sphccrocystis Schroeteri Chod. A and B, from the plankton of Loch
;

Shin, Sutherland, x450; C, zoogonidium, after Chodat ( x about 700).

methods of multiplication are :

1st, by division of the cells into
four daughter-cells, either in one plane or tetrahedrally disposed ;

2nd, by the rupture of the walls of a sporangium containing a

number of daughter-cells, with a slight retention of the old walls
of the mother-cells as in Schizochlamys ; 3rd, by the formation of
biciliated macro- and microzoogonidia. This genus very closely
resembles Gloeocystis Nag.

Sph. Schroeteri Chod. is an abundant Alga in the freshwater plankton,

occurring in all the larger lakes of the British Islands diam. of colonies 50
;

1200 /LI; diam. of cells 6 10 p.; fig. 110. Chodat regards both Tetraspora
lacustris Lemm. and Botryococcus sudeticus Lernm. as stages of this plant.
With regard to the former he is possibly correct, but the latter is a close rela-
tive of Botryococcus Braunii and certainly has no affinity with Xphcerocystis.

1
Chodat in Bull. Herb. Boissier, 1897, p. 292, t. ix.
 Palmellacece 243

Sub-family II. TETRASPORE.E.

The usually grouped in fours, or more rarely irregu-
cells are

larly disposed, towards the periphery of a structureless mass of

jelly. The sub-family is distinguished from the Palmelleae and
'

the Gloeocystideae by the pseudocilia which are attached in

'

pairs to each cell.

Genus Tetraspora Link, 1809. [Indus. Stapfia Chodat, 1897.]

The colonies are gelatinous, macroscopic or microscopic, indefinitely
expanded or more or less intestiniform and convoluted. The cells
are spherical, distributed without order near the periphery of the

enveloping jelly or grouped in twos and fours. In structure the

,
Og go
Oo o O

o
% 00
o
00
o a
>

"

Fig. 111. Tetraspora lubrica (Roth) Ag., from near St Just, Cornwall. A, uat.
size; B, portion of colony, x 100 C, x450; D, zoogonidium, x 4-".n.
;

cells are similar to those of Palinella, with the addition of t\\o

'

(or more
rarely four) pseudocilia.' Multiplication takes place
by repeated division of the cells, chiefly in two directions in one
plane, with the gelatinization of the mother-cell- walls. Reproduc-
tion occurs biciliated zoogonidia arid isogamous planogainetes.
by
Hypnospores, with thick cell-walls of a brown colour, are also

produced.
Several 'species' of this genus occur in the stagnant waters of the British
Islands, but it is very doubtful if they are specifically distinct. T. ydatinosa

(Vauch.) Desv. is the commonest form, with a vesicular gelatinous colony and
6 12 p in diameter.
cells T. lubrica Ag. (fig. Ill) and T. e.rf>l<tji>tt<i Ag. are
most probably other forms of this species. T. lacustris Lemm., which is a
form confined to the plankton, is possibly a stage in the life-history <>f
*j>

cystis Schroeteri Chod.

162
244 Chlorophycece

Genus Apiocystis Nag., 1849. The gelatinous colonies are

relatively small and attenuated
towards the base, which is usually
fixed to other larger Algse. The cells are similar in structure to
those of Palmella, and they are disposed without order near the
Each cell is furnished with
periphery of the gelatinous vesicle.
two 'pseudocilia/ which
the
penetrate through
jelly into the surround-
ing water. Multiplication
occurs by the division of
the cells in two or three
directions accompanied
in-
by a corresponding
crease in the size of the
1

colony. Correns states

that when a cell divides
one pseudocilium goes
to each daughter-cell, a
second one being subse-
quently developed. Re-
production takes place
by biciliated zoogonidia
and isogamous gametes 2 .

A. Brauniana Nag. is not

Fig. 112. Apiocystis Brauniana Nag.,

from uncommon in ponds, ditches,
near Bowness, Westmoreland ( x 400). and its pyri-
bog-pools, etc.,
forin colony is commonly
1000 M
attached by the base to larger filamentous Algse. The colonies
are 12

(or upwards) in length

and the cells 68
/x
in diameter fig. 112.
;

Sub-family III. GLCEOCYSTIDE.E.

colonies of this sub-family consist of an aggregate of

cells
The
within a common mucilaginous envelope, which exhibits a lamellose

structure. As a rule concentric coats of mucus can be distinguished

round each individual cellor round a small group of daughter-
cells. The lamellation of the mucous integument reaches a

i
Vide Bot. Centralbl. liv, 1893, p. 146.
-
Moore in Journ. Linn. Soc. Bot. xxv, 181)0.
 Palmellacete 245

maximum in some species of Glceocystis. Multiplication occurs

principally by a tetrahedric division of the mother-cells, accom-
panied by a gelatinization of the mother-cell-walls. Reproduction
also takes place by biciliated zoogonidia.

Genus Gloeocystis Nag., 1849. [Chlorococcum Fries, 1825 (in

part).] The plants occur as small irregular colonies consisting of

fsF?358

'

- -

Fig. 113. A E, Glceoci/stis infusionum (Schrank) W. & G. S. West, from Barnes

Common, Surrey. F H, Gl. gigas (Kiitz.) Lagerh., from Wimbledon Common,
Surrey. (All x420.)

an indefinite group of cells formed by the successive division

of the mother-cells. The enveloping mucus generally shows a
246 Chlorophycece
marked lamellation, similar to that exhibited by Glceocapsa among
the MyxophyceaB. The cells are globose or ellipsoid, with a parietal
bell-shaped chloroplast furnished with one pyrenoid.
The most abundant species is Gl. gigas (Kiitz.) Lagerh. [
= Chlorococcum
Gloeocystis ampla (Kiitz.) Kabenh.], which is found
Grim. in stagnant
ffigas ;

waters among other Algse; the cells are globose and 10 17 /* in diameter;
fig. 113
F H. G. vesiculosa Nag. is also a very common species of the genus
in which the cells are ellipsoid and 4 12 p in diameter. Gl. infusionum

(Schrank) W. & G. S. West \_-CJilorococcum infusionum (Schrank) Menegh.]

isa large species in which the lamellation of the integument is most remark-
able diam. of cells 25 30 p.
;
diam. of integuments of a single cell often
;

180 p; fig. 113 A E. There are several other so-called 'species' of this
genus, but they are of doubtful value.
The genus Capswlococcus Bennett (1888) is of very doubtful character and
possibly does not belong to the Algae. C. crateriformis Benn. was described
from the English Lake District.

Genus Dactylothece Lagerh., 1883 1

. This genus closely
resembles Gloeocystis in
the general arrangement
of the colony, but the
cells are oblong-ellipsoid
1

Q,
and division only takes
v
Vj A B c D place in one direction.
The chloroplast is a pari-
Fig. Dactylothece Braunii
114. Lagerh., etal plate only occupying
A, from near Bradford, W. Yorks.; B D, from , ,

near Senens, Cornwall ( x 420).

tw t uj r d o f the
cell and destitute of a
pyrenoid. The lamellation of the integuments is frequently in-
distinct.

D. Braunii Lagerh. occurs in damp situations and also in stagnant pools.

The cells are 7'5 10'5 ^ in length and 3'5 4/8 //
in breadth ; fig. 114.

Genus Palmodictyon Kiitz., 1845. The colonies of this genus

are very remarkable, the groups of cells and their surrounding

integuments being arranged in cylindrical masses, which branch

and anastomose with each other. The external mucous coats of
these elongated colonies often become very tough and assume a
reddish-brown colour. Reproduction occurs by the formation of
resting-spores or hypnospores with brown cell- walls. The germi-
1
Lagerheini in Ofvers. af K. Sv. Vet.-Akad. Forh. 1883, no. 2, t. 1, f. 2224.
 Palniellacece 247

nation of these resting-spores results almost immediately in the

formation of an elongated colony.

Palmodictyon viride Kiitz. is a very rare British Alga which I have only
observed from the extreme south-west of England. The cells are globose and
5'5 9 p. in diameter the cylindrical colonies vary from 28 52 p, in diameter
; ;

fig. 115.

'-

r
%

, -
^

Fig. 115. Falmodictyun viride Kiitz., from near St Just, Cornwall ( x420).
A, part of adult, branched colony; B and C, young colonies.

Genus Botrydina Breb., 1839. The colonies consist of sub-

spherical aggregates of cells of the Gloeocystis-iype, enveloped in
a thick gelatinous integument which may reach a diameter of
500 p. The genus requires further investigation.
B. vulgaris Breb. occurs amongst mosses on damp ground, on the trunks
of trees, etc. The cells are globose or ellipsoid and 2 7 JJL in diameter. It
sometimes occurs in large quantity amongst Leucobryum glaticnm.
 Class 4. HETEROKONTJE.
THIS class has been recently established by Luther to embrace
a number of Algae which were for a long time regarded as members
of the Chlorophycese.
are yellow-green in. colour owing to the presence of a
They
large quantity of xanthophyll in the chromatophores, and the
product of metabolism is an oil.
The class was instituted to include two series of organisms, one
'
of which, the Chloromonadales,' is
distinctly a group of the
Flagellata, including certain primitive Flagellate organisms from
which the rest of the Heterokontas have been evolved. Bohlin,
and also Blackman and Tansley, place the genus Vaucheria in this
'
class under a third series, the Vaucheriales'; but I have preferred
to retain this genus in the Chlorophycese in the old family
Vaucheriacese of the Siphonese. Excluding Vaucheria and the
Flagellate group Chloromonadales, the rest of the Heterokontse fall
'

in 1889, and they

'
under the order Confervales proposed by Borzi
are all strictly algal in organization. They are unicellular, multi-
cellular, or coenocytic in character, exhibiting a considerable

variety of form, and occurring as simple rounded unicells, long

multicellular filaments, or large gregarious cosnocytes. The cells
usually contain many chromatophores (rarely only one), which are
discoidal in form, of a yellow-green colour, and devoid of pyrenoids
and starch. In other respects the cell-structure is similar to that
described for the Chlorophyceae.
Multiplication by cell-division does not take place in some
genera.
The usual method of asexual reproduction is by zoogonidia of a
somewhat peculiar character. They are generally ovoid or pear-
shaped and are furnished with two cilia. The latter have only
been accurately examined in a few genera and have been found
to be of unequal length. They are attached to one side of the
 Heterokontce 249

anterior extremity of the zoogonidium and are generally carried in

opposite directions. Each zoogonidium usually possesses several
parietal chromatophores (vide fig.
121 C).
in each
Aplanospores are also frequently formed, usually one
cell and possessed of thick cell-walls.

Sexual reproduction occurs by the fusion of isogamous piano-

gametes which most probably resemble the zoogonidia in the
1

possession of two unequal cilia It is only very recently (1898)

.

that the second short cilium was demonstrated, up to that time

the zoogonidia and gametes of these Algae having been described
as possessing only one cilium. The second short cilium is usually
carried in a backward direction, pressed closely against the body of
the cell.

The Heterokontse contains only a limited number of genera.

None of the Flagellate series 'Chloromonadales' have been observed
from the British Islands, and all the remaining genera belong to
the Confer vales. Chlorobotrys Bohlin, which that author referred
to the Chloromonadales, is strictly algal in character and belongs
to the Tribonemaceae.

Order I. CONFERVALES.
In this order the cells are entirely algal in character, thus
differing from those of the Flagellate order Chloromonadales.
The plants are unicellular, mttlticellular, or coenocytic, and the
cell-walls are sometimes very thick. Each cell contains several or

many discoidal chromatophores, with a parietal disposition, and

from which pyrenoids are absent.
The reproduction is by zoogonidia and isogamous planogametes,
the former possessing a pair of unequal cilia.

The order is divided into three families :

Family 1. Unicellular or colonial; cells small,

Chlorotheciacece.
often attached by basal stalks, uninucleate, with one or many chro-
matophores.
Family 2. Tribonemacece. Unicellular or filamentous ;
cells often

elongate, uninucleate or subcoenocytio, with several or many chromato-

phores cell- walls rirm and thick.
;

3.
Family Botrydiacece. Plant-body large and ccenocytic, fixed,
with well-developed organs of attachment chromatophores numerous.
;

1
Gametes have been described in certain genera with two equal cilia, but such
statements are not established facts and require re-investigation.
250 Heterokontce

Family 1. CHLOROTHECIACE^.
In this family the plants are very small, unicellular, gregarious,
or colonial. The solitary and gregarious cells are attached each by
a basal stalk, generally to some larger filamentous Alga, but in
the colonial forms the cells are united by mucilaginous bands or
stalks. Each cell contains a single nucleus and one or many
parietal chromatophores. Multiplication takes place in some
genera by vegetative division, but not in others. Reproduction
occurs by zoogonidia and isogamous gametes, with one long cilium
and most probably a shorter one, the presence of which has not yet
been ascertained.
The three following genera are known as British :

A. Cells stalked ;
no vegetative division.
* Cells
gregarious, living in the mucilaginous invest-
ments of other Algse and attached by a stalk of
extreme tenuity with one chromatophore
; Stipitococcus.
** Cells ovoid, rounded, or ellipsoid, attached by a thick
stalk chromatophores several or many
; Characiopsis.
B. Cells united to form branched colonies by tubular
stalks of mucilage vegetative division abundant... Jfischococcus.
;

Genus Stipitococcus West & G. S. West, 1S98 1 . The cells are

very minute, gregarious, and

epiphytic on other filamentous
Algse, being embedded in the
mucilage surrounding the fila-
ment, to which they are attached
by long stalks of extreme fine-
ness. In shape the cells are
ovoid and apiculate, or somewhat
campanulate, with a rounded
base and an attenuated or ir-

regularly expanded apex. There

Fig. 116. Stipitococcus urceolatm
W. & G. S. West A C, epiphytic on is a single parietal chloroplast of
;

a filament of Mour/eotia, from Oughter- a pale green colour, somewhat

shaw Tarn, W. Yorks. A, x500; B ;

and C, x 780; D, epiphytic on Sphiero- irregular in form and curled

zosma excavatnm, from Harris, Outer round the inner surface of the
Hebrides (x500).
cell- wall. A single small nucleus
is present in the centre of the cell .
Reproduction is
by zoogonidia,
1
West & G. S. West in Journ. Bot. Sept. 1898.
 Chlorotheciacece 251

two of which are produced from each mother-cell. Only one long
cilium has been observed on the zoogonidia, but it is
very probable
that another shorter one has been overlooked. The zoogonidia
become attached by the end of the long cilium, the latter then

forming the stalk of the new unicellular plant.

The genus closely resembles Peroniella Gobi 1 which is also an ,

epiphyte, occurring the mucous investments of Hyalotheca.

in
The cells are, however, smaller than in that genus and their apices
are acute or expanded. The general form of the cells is thus
different from the rounded cells of Peroniella.

8. urceolatm West & G. S. "West is known from W. Yorkshire and from

N. W. Scotland, occurring as an epiphyte on Mougeotia and Hphivrozosma;
diam. of cells 3 4'2 /x; length 6'5 10'5 p; length of hyaline stalk 4 6 /*;
fig. 116.
Schinidle 2 has described another species from Germany.

Genus Characiopsis
Borzi, 1895. Most of the plants of this
genus were at one time described as species of Characium A. Br.
The cells are rounded, ellipsoid, or ovoid,
sometimes acuminate at the apex, with a
firm cell-wall and a short basal stalk of
some thickness. They occur as epiphytes,

solitary and there is no

or gregarious,

vegetative division. Each cell contains

several or many small, parietal chroma-

tophores of a pale-green colour. Repro- ^
B C
duction occurs by the division of the cell-
Fig. 117. A,
contents either to form eight (or more?) miiiutd (A. Br.) Borzi,
from near Penzance, Corn-
zoogonidia, which are set free by the
disso-
wall. B D, Ch. turtjida
lution of the upper portion of the mother- W. & G. West, from
S.
Keigblev Moor, W. Yorks.
cell-wall, or to form a number of globose
(x500).
aplanospores which become gametangia
immediately on liberation, each producing two or four gametes.
There are a number of species, several of which are known from Britain.
Ch. minuta (A. Br.) Borzi is the most frequent; length of cells 17 18^;
breadth 5-5 p fig. 117 A. Ch. tunjidi W. & G. S. West is the largest species
;

of the genus; length of cells 36 46 p breadth 1T5 ;

16 p.; fig. 117 B D.

Genus Mischococcus Nag., 1849. The cells are globular and

are united by thick tubular stalks of mucilage to form small

1
Gobi in Scripta Botan. Horti Univers. Imp. Petropolitanre, torn, i, 1866-7,
pp. 244250, t. 1.
3
Schmidle in Hedwigia, Bd xli, 1902, Heft 4, p. 153, fig. A 1.
252 Heterokontce

branched colonies ; they are situated only at the extremities of the

mucous tubes and each possesses from one to four
chromatophores.
Reproduction takes place by zoogonidia and isogamous piano-
gametes. The zoogonidia usually germinate directly to form a
small typical colony, but the
zygospore divides in two directions
in one plane
forming an epiphytic cushion, all the cells of which
are situated on short, broad,
mucilaginous stalks.

Fig. 118. Mischococcus confervicola Nag. A, from Cam Fell, W. Yorks.;

B and C, from Ruislip Eeservoir, Middlesex ( x 500).

M. confervicola Nag. is a rather uncommon Alga, occurring as an

epiphyte
on various filamentous species of Cladophoracese,
Tribonemacete, Ulotrichacete,
(Edogoniacese, etc. It is generally found in small and
ponds ditches, or more
rarely in peaty pools; diam. of cells 3'5 5'5 /*; fig. 118.

Another genus belonging to the Chlorotheciaceae has been

recently described by Schmidle under the name of Oodesmus.
1

Lemmermann reports having found this genus in some plankton

material forwarded to him from Loch Doon, The cells
Ayrshire.
are ovoid and united by short bands of mucus to form free-

swimming colonies. Each colony consists of four cells, which are

disposed in one plane. The cell-wall is relatively thick and there
are one or two chromatophores in each cell. 0. Doederleinii
Schmidle is the only known species cells 8 p in and 6 in
;
length //,

breadth.

1
Schmidle in Hedwigia, Bd xli, Heft 4, 1902, p. 162, fig. B 4.
 Tribouemacece 253

Family 2. TRIBONEMACE^E.
The
plants of this family are unicellular or filamentous. The
cells are globose, cylindrical, elongate, often spirally coiled or
united to form long flexuose filaments. There is generally a single
nucleus in each cell, but sometimes two or more are present,
giving the cell a more or less ccenocytic character. The cell-walls
are always firm and usually of some thickness, except in the genus
Bumilleria, in which the walls sometimes become swollen and
hyaline. Asexual reproduction takes place by zoogonidia with one
long and one short cilium, and two or several parietal chromato-
phores. Aplanospores are also of frequent occurrence in the genus
Tribonema. Sexual reproduction occurs by isogamous piano-
gametes which have been described, but perhaps erroneously so,
as having two equal cilia.

There are four British genera, three of which are abundant.

A. Plants unicellular.
* Cells
globose, aggregated in mucilaginous colonies ... Chlorobotrys.
** Cells and often
elongate, usually shortly stipitate
spirally coiled Ophiocytium.
B. Plants filamentous.
* Cell- walls
firm, splitting into H -pieces Tribonema.
** Cell-walls
hyaline, H -pieces not very evident; fila-

ments small Bumilleria.

1
Genus Chlorobotrys Bohlin, 1902 The cells are globose or .

subglobose, solitary, or more commonly aggregated in families of

2, 4, 8, or 16. Each family is surrounded by an ample mucous

integument, very hyaline and quite homogeneous. The cell-walls
are firm, smooth, of some considerable thickness, and they contain
a certain proportion of silica. The chromatophores are parietal,
yellow-green discs, from 6 to 30 of which are disposed on the walls
of each cell. Sometimes the pigment becomes more or less diffuse.
There is frequently a prominent red pigment-spot in each cell of
the family, but sometimes this is entirely absent. Multiplication
takes place by the division of the cells primarily in two directions,
but afterwards in three directions-. Families of 4, 8, or l(j cells
are therefore frequently very symmetrical, but beyond this number

they become irregular. During cell-division the contiguous walls

1
Bohlin iii Bihaiig till K. Sv. Vet.-Akad. Handl. Bd 27, no. -i, 1902, p. 34,
t. i, f. 9.
-
West & G. S. West in Journ. Bot. April, 1903.
254 HeteroJcontce

of the daughter-cells are at first much flattened, but afterwards

become more convex. Zoogonidia have not been observed.
Bohlin was distinctly in error in referring this Alga to the

Chloromonadales, as the plants are strictly algal in character.

The ordinary vegetative
condition of the orga-
nisms belonging to the
Chloromonadales is a
ciliated or flagellated
one, whereas a motile
conditionofChlorobotrys
has never been observed.
Moreover, Chlorobotrys
isa very abundant Alga
B and one which I have
had under observation
Chlorobotrys reyiilaris (West) Bohlin
Fig. 119.
from Tremethick Moor, Cornwall ( x 450). formany years and the ;

motile condition, far

from being the ordinary state of the plant, must be exceptionally rare.
The genus is undoubtedly nearest to Botrydiopsis Borzi, but
differs in the smaller size of the cells, which are associated to form

colonies, and the prominent mucous investment. Moreover, it is

still water, whereas Botrydiopsis inhabits running
only found in
water.
Chi regularis (West) Bohlin \Chlorococcum regulare West ] is widely dis- 1

tributed and often abundant in the Sphagnum-bogs of the British Islands.

I have also examined numerous specimens of it from Norway, Switzerland
and the United States. The cells are 12 19 p in diameter and the families
with the mucous investments) are 34 90 p, in diameter; fig. 119.

Genus Ophiocytium Nag., 1849. [Indus. Sciadium A. Br.,

1855.] The cells are free or attached and generally many times
longer than the diameter. They are commonly solitary, but
sometimes colonial, and they are usually curved or spirally con-
torted. The apex of the cell may be capitate or apiculate and
is sometimes attenuated into a long spine. Each cell contains
several nuclei and a number of large parietal chromatophores of a
somewhat cylindrical form. The presence of oil drops is a feature
of some species. The cell- wall consists of a homogeneous lid fitted
to the apex of a long tube composed of apposed layers of pectose
1
West in Journ. Boy. Micr. Soc. 1892, p. 737, t. x, f. 55.
 Tribonemacece 2f>r>

compounds. Vegetative division does not occur. Asexual repro-

duction takes place by the division of the cell-contents to form
ellipsoidal aplanospores, or to form eight
ovoidal zoogonidia with
two cilia. In those
species which are at-
tached the zoogonidia
generally come to rest
on the rim of the empty
tube-like cell and there

grow into adult cells.

A repetition of this pro-

cess produces a curious

branch-system. Some-
times the zoogonidia
develop on the apices
of other cells which
contain the cell-con-
tents. No gametes have
been observed. The Fi S- 12 0- A Ophiocytium majus Nag., from
>

Bowness, Westmoreland. B G, 0. cochlrare

genus was monographed (Eichw.) A. Br., from same locality. H aud I,
O. bicuspidatum (Borge) Lemm. forma longispina
by Lemmermann but
1
,
Lemrn. from Pilmoor, N. Yorks. J, O. Arbii?.-
,

the validity of the thir- rulii (A. Br.) Babenh., from Mitcham Common,

teen species he puts Surrey. (All x 450.) K, 0. yraciliceps (A. Br.)

Babenh., after treatment with potassium hydrate
forward is questionable. (after Bohlin, x 570).
Several species are widely distributed in the British Islands. 0. Arbuscula
(A. Br.) Rabenh. is an attached species, often with very pretty branch-
systems; diam. of cells 3 8/z; fig. 120 J. 0. majus Xag. is the largest
species, the cells reaching a diameter of 17 fig. 120
A. 0. cochleare (Eichw.)
fj. ;

A. Br., 0. capitatttm Wolle, 0. l>i<-tixi>t<.l<it>tin (Borge) Lemm. and 0. parrel nn,

(Perty) A. Br. are also frequent species.
Genus Tribonema Derbes & Solier, 185G. [Conferva in the
sense used by Lagerheim, 1888.] The plants of this genus are
simple filaments of cylindrical or slightly barrel-shaped cells with
strong cell-walls. The latter are often of considerable thickness
and the filaments frequently break up into H -pieces. Each
H-piece consists of a transverse cell-wall with a cylindrical piece
on either side, and the whole is composed of a number of layers of
pectose compounds. Each cell is thus bounded by the halves of
two H -pieces. The cells contain one (or sometimes two) nuclei
and a variable number of parietal chromatophores. In some
1
Lemmermann in Hedwigia, Bd xxxviii, 1899, pp. 20 38, t. iii & iv.
256 Heterokontce

species the chromatophores

are few and irregular, but more often
they are numerous and discoidal. Asexual reproduction takes
place by the formation of globular or ellipsoidal aplanospores,
which escape by the breaking up of the filament (fig. 121 B and I);
also by zoogonidia with a pair of unequal cilia (fig. 121 C).
Sexual reproduction occurs by the conjugation of isogamous
is not strictly isogamous, as one
gametes, but the conjugation
comes to rest and rounds itself off before another swarms
gamete
up to it and conjugates with it. The structure of the cell-wall of
was well described by Bohlin who also demonstrated
1
this genus ,

the close affinity between Tribonema (Conferva Lagerh.) and

Ophiocytium.
Concerning the abandonment of the generic
name Conferva,' '

I cannot do better than quote at length the remarks recently

made by Hazen 2
. He writes:

" The name Conferva is very ancient, going back to the time of Pliny. As
a modern generic name it has received most varied treatment, and covered at
different times very diverse groups of plants.
"Under this name Linnaeus included a very large part of the branching,
as well as the simple, filamentous algae. He adopted the genus from Dillenius.
species mentioned by Linnaeus, Conferva ripularis,
is undoubtedly
The first

the oldest of his group, so far as the history of these ill-defined forms can be
determined. This species, according to the synonymy of Linnaeus (Sp. PI.
1164. 1753), is Conferva fluviatilis, sericea vulgaris et fluitans of Dillenius
this in turn is Conferva Plinii Dillen. (Cat.
(Hist. Muse. 12, pi. 2,
f. 1. 1741) ;

Plant, Gissam nascentium, 199. 1719); the earliest description of

sponte
C. Plinii that we have seen is in L'Olfel's Plantarum Observationes, 664. 1576,
but undoubtedly the name is of more ancient origin.
"Now no one would imagine that the ancient herbalists or even Linnaeus
could the numerous filamentous forms known to us only by the
distinguish
use of good microscopes. Indeed, the fact that Linnaeus described only two
unbranched species is sufficient proof of this. Conferva rivularis as collected
and at another time an
by him, was very likely at one time a Spirogyra
This however, as interpreted by the earlier
(Edogonium. type species,
algologists, e.g., Dillwyn, Lyngbye
and Mueller, is evidently a form belonging
to what is now known as Rhizodonhun, and has come down to us as R. rivu-
lr'ts (L.) Kiitz. This identification is also confirmed by Linnaeus himself,
who (Sp. PI. Ed. 14. 1784) quotes the figure of C. rivularis from Flora Danica.
"At any rate, there is no warrant whatever for employing the name Con-
fervato designate the genus recognized under that name in Lagerheim's
revision, for there is no evidence that these species were ever collected by

1
Bohlin in Bihang till K. Sv. Vet.-Akatl. Handl. 1897, Bel 23, no. 3.
-
Hazen in Memoirs Torr. Bot. Club, xi, 1902, no. 2, pp. 181183.
 Tribonemacece 2f>7

Linnajus, and certainly none of them were distinguished by him from other
simple filamentous forms.
"For Lagerheim's group of species, as for all genera, the adoption of a
generic name based on a recognizable species, as a type, is essential. The

Fig. 121. AG, Tribonema bombycinum (Ag.) Derb. & Sol.; A, from Shipley,
W. Yorks. B, showing aplanospores (ap), from Senens, Cornwall; C, zoogo-
;

nidium, and D F, young plants, from Senens, Cornwall (all x 450) G, after ;

treatment with potassium hydrate (after Bohlin, x 570). H and I, T. bomby-

cinum forma minor (Wille) nob.; H, from Shipley, W. Yorks.; I, showing
aplanospores (ap), from near St Just, Cornwall (x450). J, Bumilleria pumila
W. & G. S. West, from near Senens, Cornwall ( x 450).

earliestsuch name in the present case is Tribonema Derbes & Solier (1856) 1 .

This genus was based on a single species, Conferva bombycina, and in the
diagnosis, for the first time in the history of the species, explicit mention was
made of the most essential character, namel}-, the form of the chromatophores,
although it had been previously suggested in the plates of Kiitzing. The
method of zoospore dispersal was also first described and illustrated by Derbes
& Solier. There is, then, every reason for employing the name Tribonema as
a memorial of the discernment of these authors."
1
Derbes & Solier, Me"m. sur quelques points de la physiologic des Algues, 1856.

W. A. 17
258 HeteroTiontce

T. bombycinum (Ag.) Derb. & Sol. ( Conferva bombycina Ag.) is general

throughout the British Islands; diam. of cells 8 15 p.; fig. 121 A G. A
small form of this species [forma minor (Wille) nob.J is also very common;
diam. of cells 5 6'5 p, fig. 121 H and I.
: T. affine (Ku'tz.) nob. (^Conferva,

affinis. Klitz.) is the thinnest species with the most elongate cells, and the

chromatophores are few and irregular; diam. of cells 5 5'4 p.. T. obso-
letum nob. ( = Conferva obsoleta West & G. S. West) is a much rarer species ;

diam. of cells 19 21 p..

Genus Bumilleria Borzi, 1895. The cells of this genus are

arranged in long filaments which differ from those of Conferva

principally in the structure of the cell-wall. The latter is

practically homogeneous and does not readily break into H -pieces.

The pectose constituents instead of forming closely apposed layers
may form a distinct mucous cylinder in which the cells appear to
be embedded. The chromatophores are small, pulvinate and
parietal.
B. pumila West & G. S. West is the only known British species ; length of
cells 56 p.; diam. 4'8 5'7 p.; fig. 121 J.

Family 3. BOTRYDIACE^.
This family is well marked off from the other groups of the
Confervales by the form of the plants alone. Each individual is
a rounded coenocyte of macroscopic size and is attached to damp
mud by well-developed rhizoids.'
'
The chromatophores are very
numerous and the methods of reproduction are somewhat
diversified.
Genus Botrydium Wallroth, 1815. The plants consist of

green, pear-shaped or spherical coenocytes of considerable size,

growing on damp mud into which they are rooted by a branched

system of colourless rhizoids. The coenocytes are vesicular, with a
lining layer of protoplasm in which are embedded numerous nuclei
and chromatophores. The latter are irregular in form, more or less
evenly scattered in one or more layers through the lining proto-
plasm, and are generally in close contact with the nuclei. Bodies
of the nature of pyrenoids have been observed in the younger

stages of the plant, but starch is not formed. The rhizoids possess
protoplasmic contents and many nuclei. Wager has observed
mitotic division of the nuclei and states that the chromatic sub-
stance appears to reside wholly in the nucleolus.
This plant reproduces itself asexually in a great variety of
ways, depending largely on the 'conditions of environment, any
 BotrydiaceoB 259

change of conditions usually resulting in a corresponding variation

in the reproductive process. These different processes have been
worked out by Rostafinski
and Woronin and the 1

final result is in each case

either the production of

zoogonidia or aplano-
spores. The zoogonidia
are and ovoid in
small
shape, with one or two
chromatophores and one
long cilium. (Very pro-
bably there is a second
shorter cilium, but its

presence has not yet been

ascertained.) The whole
plant frequently becomes
a huge zoogonidangium,
it becomes
especially if
submerged, and the zoo-
gonidia escape through
Fig. 122. Botry ilium granulation (L.) Grev.,
an apical opening. The from Calverley, W. Yorks. A, nat. size; B and
C, x45; D and E, x450; D, aplanospores; E,
aplanospores, which are
germinating aplanospore.
globose or ellipsoid, often
become hypnospores, and they are frequently produced in numbers
in the rhizoids. On
the green portion of the plant above ground
becoming too dry, the contents migrate into the rhizoids and a
large number of aplanospores are formed. The development of the
young plants much, depending upon the external conditions.
varies
Woronin described a sexual reproduction by
Rostafinski and
the conjugation of isogamous gametes, but Klebs has given good
reasons for doubting this.
B. yranulatum (L.) Grev. is a very local plant. It occurs widely distributed
over the British Islands, but the conditions are not often suitable for its
appearance above ground. It is found almost exclusively on drying-up mud,
and sometimes occurs in countless numbers on mud turned out from a canal
or on the drying bottom of a muddy pond. I have observed these plants so

thick as to stand out in mulberry-like masses from the surface of the damp
mud. The nature of the mud is immaterial and the Alga is not uncommonly
found on chalk mud. The plants reach a diameter of 2'25 mm. Fig. 122.

1
Rostatiuski & Worouiu in Botan. Zeitung, xxxv, 1877.

172
 Class 5. BACILLARIE^.
THIS class of Algse includes a large number of minute plants
known as the Diatoms. They are perhaps better known under '

of the Diatomacese, but the earlier name Bacillarieae

'

the name
has been in use for many years among systematists, particularly
'

and the name Diatomacese is here retained

'

in continental Europe,
for one family only. The class is a very large one, with well-
marked characters, and includes about 10,000 species. As would
be expected among such a large number of species there is great
variability of form, but
at the same time the Diatoms always

possess those salient features which mark them off from all other
Algse.
They are universally distributed in both fresh and salt water,
and as the wonderful sculpture of their cell-walls renders them
objects of great beauty, they
have long been made the subject of
special study by
numerous students of natural history.
Diatoms are unicellular plants, mostly of minute size, the cell-
walls of which are composed of an organic matrix impregnated
with silica. The silica can be removed by the action of hydro-
fluoric acid, leaving the organic matrix behind or the organic ;

matrix, which is allied in composition to cellulose, can be. removed

by calcination leaving behind the siliceous constituent.
Each individual Diatom is termed a frustule, and the cell-wall
consists of two more or less equal valves, joined together by two
connecting -bands which overlap. Each half of the Diat* i is thus
and the
composed of two pieces, a valve and a connecting-band,
that of the younger
connecting-band of the older half fits over
half like the lid of a cardboard-box. The connecting-bands,
are distinct from
although closely fitted to their respective valves,
them, and the two bands together form what is termed the f/irdle.
The latter does not usually consist of two closed hoops, but as
 Eacillariece 201

Palmer and Keeley 1 have pointed out, each band of the girdle is a
two-ended band of silica with the ends overlapping without being
joined. Each frustule possesses a thin coat of mucus which can
be readily demonstrated by slight staining with aniline dyes.
Diatoms often occur as solitary, free-floating individuals, but
they may adhere to one another to form chains, either by their
valve-faces to form ribbon-like or
thread-like colonies (Ewnotia, Melo-
sira), or by mucous cushions at
their angles to form zig-zag colonies

(Tabellaria). Some adhere closely

to larger plants by the whole of one
valve-face (Cocconeis), or they are
affixed to some object by
larger
simple or branched gelatinous stalks
(Gomphonema); others occur in large
colonies embedded in a common
mucilaginous envelope, either as a
compact mass or a simple or branch-
ed tube. This condition is commoner
in marine species than in freshwater

ones, and is variable even in indi-

IV
vidual species. Marine Diatoms
also reach a much larger size than
freshwater ones.
In referring to any individual Fig. 123. A, valve view of
Navicula nobilis Ehrenb. var. Dac-
Diatom, the aspect in which the ti/lnx (Ehrenb.) V. H., from Dol-
Wales x 250).
girdle side is exposed to view is gelly, ( B, girdle
view of major Kiitz., from
N.
best termed the
girdle-view (or Shipley, W. Yorks. (x300). en,
central nodule iv, inner valve
zonal-view), and that in which the ; ;

ov, outer valve; pn, polar nodule;

surface of the valve is exposed to r, raphe.
view the valve-view.
The valves are as a rule thin and transparent, slightly convex
on the c.ftside, and in almost all species they are ornamented with

variously disposed strife. The best lenses, however, have shown

that these striae consist of series of small cavities within the
siliceous wall of the Diatom, and it is their close and regular
arrangement that causes them to appear as striae. The striae are

Palmer and Keeley in Proc. Acad. Nat. Sci. Philadelphia, 1900.

2G2 Badllariece

so extremely fine and constant in some species as to furnish

splendid test-objects for the definition and angular aperture of the
lenses of microscopes. The valves of some genera, particularly the
marine ones, exhibit a beautiful areolated structure, due to the
presence of chambers in the siliceous cell-wall. These chambers
may be open to the exterior or covered by a thin membrane, and
their inner walls are perforated by exceedingly minute apertures
or pores which lead into the cell. These pores are not present in
all species of Diatoms, Miiller and Lauterborn having shown that
in some species of Navicula (Pinnidaria) they are probably absent.
Schlitt affirms that there is no question of the existence of pores
in many species of Diatoms, and their existence in a large number
of others is extremely probable. He draws a distinction between
pores and dots, and this has been further emphasized by Miiller.
The latter has termed small circular dots which resemble pores
1

'poroids'; he gives O'lyu, as the minimum diameter of pores, and

0'4 0'5/i as their maximum diameter, and all structures over
3
O'Gyu, he regards as Miiller recommends treatment
2 '

poroids .'

with hot sodium carbonate and potassium hydrate in studying the

structure of the cell-wall.
Heribaud 4 states that increased altitude and enfeebled light
cause a diminution of the number of striae and of their strength,
accompanied by an increase in the length and breadth of the valves.
Many Diatoms exhibit a thickening of the cell-wall, visible in
the valve- view, in the centre of the valve and very often at both
extremities. These thickenings are known as nodules. The
nodules are very frequently connected by a long median line
known as the raphe. If the central nodule spreads out in a
lateral direction it is known as a stauros. A portion of the valve
on each side of the raphe and round the central nodule is often
quite devoid of stria? this plain or smooth portion of the valve is
;

sometimes spoken of as a hyaline area.

The raphe, for at least some part of its length, is a true cleft
in the valve through which the protoplasmic contents of the cell
are placed in communication with the surrounding medium.
(Consult Fig. 124r.)

1
0. Miiller in Berichte Deutsch. Bot. Gesellsch. xvii, 1899.
-
O. Miiller, torn. cit. xviii, 1900.
3
0. Miiller, torn. cit. xix, 1901.
4
Heribaud in Comptes Bendus, cxviii, 1894.
 Badllariece 203

Some Diatoms possess a pseudo-raphe, which is a simple line,

central or sub-central, and without a central nodule. Often the
conspicuous in both valves, but sometimes
is it is
raphe only
present in one valve (Achnan-
tJies, Cocconeis, Rhoicospkenia).
In many Diatoms the raphe is
obscure and marginal or sub-
marginal. In some species, such
as Navicula nobilis'Eihrenb., there
is a smaller accessory raphe which
lies alongside the primary one,
and is united to it close to the
central and terminal nodules.
In Vanheurckia there is a sili-
ceous rib on each side of the
Fig. 124. Transverse section of the
raphe and nodules, and in Am- frustule of a Navicula. (From Van
Heurck, after W. Prinz.) cb and cb',
pin'pleura the central nodule is
the two connecting-bands forming the
girdle; r, raphe; en, central nodule;
greatly elongated.
c, costa of valve. (Very highly magni-
Some Diatoms (Fragilari- fied.)
oidese,Mastogloia) possess longi-
tudinal septa which are always more or less considerably perforated.
These septa are parallel to the valve-faces and are usually situated
between the girdle and the valves.
Each individual Diatom contains a more or less centrally dis-

posed nucleus, generally with a nucleolus. A very weak solution

of inethylene blue will bring out the nucleus in living Diatoms,

staining it clearly before the rest of the protoplasm. The proto-

plasmic mass surrounding the nucleus is connected with the
primordial utricle either by two broad bands of protoplasm or by
a number of radiating or anastomosing threads. One or several
large vacuoles occupy the central portions of
the cell.
The chromatophoresof Diatoms are somewhat variable. One
or many may be present in each cell; they may be small and
discoidal, large and plate-like, or extensive anastomosing masses

occupying a large part of the lining protoplasm. In many species

they are extremely irregular in form, being band-like, lobed, or
even presenting the appearance of perforated plates. They are of
a golden-yellow or brown colour, or very rarely green, as in some
forms of Navicula viridis Kiitz. and JV. cuspidata Kiitz. They
contain chlorophyll, but this is masked by the presence of a brown
264 Bttcillai'iece
/
pigment known as diatomin, which resembles the phycophsein of
the Phseophycese. The diatomin, which can be extracted by
alcohol forming a yellow-brown solution, is itself a complex sub-
stance containing, amongst other pigments, xanthophyll. An
alcoholic solution of diatomin turns a beautiful
blue-green colour
on the addition of sulphuric acid. The chromatophores contain a
variable number
of pyrenoids which often project into the interior
of the cell as rounded elevations. 1
Mereschkowsky has observed
pyrenoids which have partially or entirely emerged from the
chromatophores, appearing as free colourless bodies on their inner
surfaces.
The nutrition of the vast majority of Diatoms is
holophytic,
but a few saprophytic forms are known'-. The latter are peculiar
in the complete absence of pigment, and they
apparently occur
in water in which there is an abundance of
decaying organic
matter 3 Karsten 4 has found that Nitzschia palea (Kiitz.) W.Sm.
.

when cultivated in favourable nutritive media will become

saprophytic.
Diatoms are incapable of growth in size owing to the siliceous
nature of their cell-walls, but slight alterations of volume can take
place by a sliding movement of the connecting-band of the older
half of the cell over that of the younger half.
In addition to the symmetrically arranged markings on the
valves, the frustules of Diatoms possess an external symmetry in
one or more planes. Some
of them are zygomorphic in one plane
only, some in three planes at right angles, and others exhibit a
radial symmetry.
The movements of Diatoms Most of the solitary, unattached
:

species of Diatoms exhibit movements which have long been a

puzzle to students of
biology. This power of locomotion is
especially in marked
species of a naviculoid form, and various
explanations have at different times been put forward to account
for it. The movements of some forms are very slow, but others
are capable of propelling themselves with considerable
rapidity
backwards and forwards in the direction of their longer axis. This
spontaneous movement is sometimes creeping and steady, but at
1
Mereschkowsky in Flora, xcii, 1903, pp. 7783.
The following
'

are colourless saprophytic Diatoms: Nitzschia putrida Benecke,

N. leucosigma Benecke, and Synedra hyalina Provasck.
3
Benecke in Pringsheim's Jahrb. f. wissensch. Bot. xxxv, 1900.
4
Karsten in Flora, Ixxxix, 1901.
 Bacillariece 205

other times jerky, and is generally along a fixed substance with

which the Diatom is in contact. It is not at all comparable to
the free-swimming movements of many other Algte, arid, as a rule,
only takes place when one valve-face of the Diatom is in contact
with the fixed object.
Ehrenberg (1838) imagined the movement to be due to the
of
protrusion of cilia or of a pseudopodium through the raphe
the valve, whereas Nageli (1849) attributed it to the passage of
osmotic currents through the cell-wall. Max Schultze (1865)
1
,

who observed the movements of minute foreign particles down

the length of the raphe, attributed the locomotion to the contrac-
tility ofa small portion of the protoplasm which was protruded
through the raphe. Hallier (1880) considered it to be due to
a contractile layer of protoplasm, and Onderdonk (1885) also
regarded it as due to an external movement of protoplasm, but
2
Mereschkowsky (1880) concluded that the evidence was in favour
of Nageli's theory of osmotic currents through the cell-wall.
3
0. Miiller (1889) demonstrated the presence of a large number of
minute pores and anastomosing fissures in the valves of certain of
the large species of Navicula (Pinnutaria), and showed that the
central and terminal nodules are traversed by straight and curved
which run towards the raphe and are eventually merged
canaliculi
with Owing to intracellular pressure the protoplasm emerges
it.

from the pores of the central or terminal nodules and passes

down the whole length of the raphe, returning into the cell-cavity
through the pores of the terminal or central nodules as the case
may be. There are thus two sets of currents on each valve of the
Diatom. The cause of the movement of the frustules was ascribed
by Mtiller to the reaction of the motive forces of this living stream
4
of protoplasm upon the surrounding water. Schiberszky (1891 ) ,

from observations on Synedra, agreed with Pfitzer that the move-

ment was due to a of protoplasm which escapes from the
coating
raphe, and which is in a condition of
vibratile motion. He believed
that the currents along the raphe were usually interrupted jerking
or pulsating movements.
Cox (1890) revived the idea
5
of a line of cilia along the raphe,

1
Max Schultze in Archiv fiir Mikr. Anat. Bd
i, 1865, p. 376.
2
Mereschkowsky iu Bot. Zeitung, 1880, 529.
p.
3
O. Miiller in Berichte Deutsch. Bot. Gesellsch. Bd vii, 1889.
4
Schilberszky in Hedwigia, xxx, 1891.
5 J. D. Cox in The Microscope, July, 1890.
266 BwillarieoB

and suggested that the absence of silica along this line could be
accounted for by the obstruction of the moving cilia. Btitschli (1892)
also imagined that the presence of a cilium or a fine flagellum
would explain the phenomenon, but no methods of staining have
ever demonstrated the existence of such structures.
The movements of some of the larger species of Navicula
(Pinnularia) have been explained by Biitschli (1892) and by
1

Lauterborn (1894)- to be due to the production of a delicate

filament which is protruded from the raphe at a point opposite
the central nodule. The frustules of Navicula major Ktitz.,
N. nobilis Ehrenb., and N. viridis Klitz. are surrounded by a
distinct mucilaginous envelope, and the protruded filament is

quite colourless and transparent, resisting all attempts to stain

it with aniline
dyes. It lies alongside -the raphe, but not in
contact with it, and it elongates by a series of jerks. Biitschli

puts this forward as the explanation of the jerky movement of

Diatoms, the frustule being pushed backward by the elongation of
the filament, the distal end of which is fixed to the substratum.
In 1893 O. M tiller 3
again emphasized his previous explanation
of the of Diatoms, affirming that they were dependent
movements
on the forces connected with the protoplasmic currents on the
surface of the valve, and he denied that the movements could be
the result of the filaments described by Biitschli and Lauterborn.
He likewise stated that these filaments were composed of proto-
plasm, and not of mucilage. Lauterborn (1894) 4 contested that
the production of motility by the streaming of protoplasmic
currents, as suggested by Miiller, would be an isolated phenomenon
in either the vegetable or animal kingdom, whereas movements
are known to occur in the Desmidiacesfi and Oscillatoriacese as a
result of the excretion of mucilage, and, according to Schewiakoff,
in the creeping Gregarinidse also. Miiller (1894) 5 replied again to
the criticism of his hypothesis, and stated that the analogy which
had been drawn between the movements of Diatoms and of
Desmids was a false one.
There is no doubt that of the smaller motile species
in many
there is a complete absence of gelatinous filaments such as those
1
Bh'tschli in Abhandl. naturh.-med. Ver. Heidelberg, 1892, Bd iv, Heft 5.
2
Lauterborn in Bericbte Deutsch. Bot. Gesellsch. Bd xii, 1894, p. 73.
a
O. Miiller in Berichte Deutsch. Bot. Gesellsch. Bd xi, 1893.
4
Lauterborn, torn. cit. xii, 1894.
5
0. Miiller, torn. cit. xii, 1894.
 Baeittariece 207

occurring in the larger species of Navicula (Pinnularia),

even
though the movements of the frustules, the structure of the valves,
the system of fissures and pores, and the arrangements of the
protoplasmic currents are the same. In some of the small species
of Navicula the movements are extremely active, comparing not

unfavourably with the slow movements of certain of the Infusoria.

If the movements of such Diatoms be clue to the secretion of
gelatinous material, then the amount secreted in a short space of
time must be relatively enormous.
After a careful consideration of the facts which have been
brought forward with a view to the elucidation of this most
interesting problem, there appears to be no doubt that the move-
ments are connected with the raphe, and the balance of evidence
indicates that in some Diatoms at least they are due to an exuda-
tion of mucilage.
The multiplication of Diatoms takes place by successive
bipartitions, each division resulting in a gradual reduction in the
size of the individuals. A slight increase in the volume of the
frustule is the first appreciable change, after which the nucleus
divides karyokinetically. A
division of the cell-protoplasm now
takes place, and a new siliceous valve is formed over each divided
surface. These new valves are situated within the girdle of the
original frustule,and the connecting-bands of the new valves are
soon developed, sometimes making their appearance before and
sometimes after the separation of the individuals. Thus, each
individual consists of a new valve and an old one, the connecting-
band of the old valve overlapping that of the new valve. Some-
times the cells do not separate, but remain in contact after division,
successive bipartitions resulting in a chain of individuals. Owing
to the formation of a pair of new valves within the girdle of the
old ones, and since the cells when once formed are incapable of

growth, every succeeding generation becomes reduced in size by

the double thickness of a connecting-band. This statement is
not strictly true, however, in the case of some of the filamentous
species, and possibly in many others. It has been shown that
daughter-cells are often produced of larger size than the parent-
cells, such daughter-cells being recognizable by the thickened rim
of the valves. This fact has a retarding influence on the diminu-
tion of the size of the cells, the reduction in size not being in
strict proportion to the number of bipartitions ; and, concerning
268 BacillariecK

1
the multiplication in the filamentous genus Melosira, Mitller
has drawn up a definite law of division. He has shown that
the multiplication of the cells takes place in such a manner
as to prevent as much as possible the division of the smallest

daughter-cells. This law, although indicating the prevailing

conditions of multiplication in Melosira, is not true of all
Diatoms.
On the greatest diminution of size having been reached for
one species, the maximum size of the species
is regained by
any
the formation of an auxospore, and there are five methods of
reproduction by auxospores.
(1) The protoplasm of a cell of the smallest size (sometimes
termed a microfrustule ') swells up and forces apart the halves
'

of the frustule, escaping to the exterior enveloped in a cellulose

membrane. This is the auxospore, the wall of which rapidly
becomes silicified and assumes the markings characteristic of the
but the form of the cell is usually very different from that
species,
of the original frustule, and often most irregular. This large,
newly-formed cell of irregular appearance almost immediately
undergoes division, the individuals of each succeeding generation
2
rapidly regaining their characteristic form and elegance. Miquel ,

who has made a special study of the manner in which the maximum
form of a Diatom is re-established, based upon experimental
cultures of a number of species, states that such re-establishment
of the maximum size is habitually brought about by the formation
of this simple type of auxospore. It is merely the rejuvenescence
of a single cell accompanied by an increase in size. (Fig. 125 C
and D.)
(2) Two auxospores may be produced by the division of the
contents of a single frustule. Each of the two portions of the
cell-contents emerges from the cell and develops as in the first
method. This method has only been observed in Rhabdonema
arciiatum (Ag.) Kiitz. and Achnanthes longipes C. Ag.
(3) An auxospore may be formed by the conjugation of the
contents of two frustules. The two Diatoms become enveloped in

common mucous covering, and the cell-contents emerge and

a fuse

to form a single body, which then develops into an auxospore.

Miiller, 'Die Zellhaut und die Gesetze der Zelltheilungsfolge von Melosira
1

arenaria Moore.' Berlin, 1883.

-
Miquel in Annal. de Micrographie, iv, 1892.
 BadllarwoB 209

This is a true conjugation of aplanogametes with the formation of

a zygospore. (Fig. 125 B.)
(4) frustules approximate and the cell-contents
Sometimes two
throw off the old valves, but there is no conjugation. The two
rounded masses of cell-contents
lie close together or separated
by some of the enveloping jelly,
and each develops independently
into an auxospore.

(5) A pair of Diatoms ap-

proximate, but before conj ugation
the protoplasm of each cell divides
into two daughter-cells. Two
auxospores are then formed by
the fusion of a daughter-cell from
each mother-cell with a daughter-
cell from the opposite mother-
cell. This is known to occur in
Amphora ovalis Klitz.,
Epithemia
Argus (Ehrenb.) and Na-
Kiitz. 1 ,

vicula limosa Ktitz. (Fig. 125 A.)

A normal auxospore can be
regarded as one produced by the
125. Naviculd
conjugation of two cells (or ga- Fig. A,
Kiitz., from the New Forest, Hants.
metes), those produced without (x450). B,Achnanthesjlexella(Eiitz.)
Breb., from Craig-an-Lochan, Scot-
conjugation being parthenoge- land (x450). C, Navicula Amphis-
netic. Theand the fourth
first bcena Bory, from Barnes Common,
Surrey (x450). A', i-iridis Kiitz.,
methods are the ones most fre-
from Clougb, Antrim, Ireland ( x 350).
quently observed. The most im- C and D illustrate the first method of
auxospore formation, B, the third
portant feature in the formation method, and A the fifth method.
of an auxospore is the increase in
size of theKarsten 2 considers that the majority of Diatoms
cell.

exhibit undoubted sexuality.

Castracane and other observers have recorded another method
of reproduction by the formation of small spores within the
frustules. Kitton 3 and Lockwood 4 have each stated that Diatoms
as to pass through
may possess spores (" micros-pores ") so small
1
Klebahn in Jahrb. fiir wissensch. Bot. xxix, 1896.
a
Karsten in Biol. Centralbl. xx, 1900; Flora, Ixxvii, 1900.
3
Kitton in Journ. Quekett Micr. Club, ser. 2, ii, 1885, p. 178.
4
Lockwood in Journ. New York Micr. Soc. 1886, ii, p. 153.
270 Bacillariecv

filter-papers, but these observations require confirmation,

as they do
1
not seem to agree with the researches of Miquel. Castracane has
stated that the normal method of reproduction of Diatoms is by

spores or germs, and that multiplication by division, although

very common, is the exception rather than the rule. This state-
ment I consider to be erroneous, as I am fully convinced from my
own observations of these plants that the normal method of increase
isa multiplication by cell-division.
Sometimes a resting condition occurs, a pair of new valves
being formed within the old ones. This corresponds to the forma-
tion of aplanospores in the Chlorophycese, and has been termed
the craticular state.
Diatoms are amongst the commonest of microscopic objects,
and they are ubiquitous in all kinds of damp and wet situations.
They occur in fresh, brackish, and salt water, often forming a
yellowish-brown scum at the surface, or thickly clothing larger
Algai- or other aquatic plants. They are most abundant in cold
latitudes, having a decided preference for cold water, although
some of them have become adapted to life in hot-springs. In the
ocean they are more abundant than any other pelagic plants, and
in the cold surface-waters of the Arctic and Antarctic Oceans they
occur in prodigious quantity. They form a considerable part of
the food of many freshwater and marine animals, and are often
found in quantity in the alimentary tracts of Molluscs, Crustacea,
Tunicates and Fishes. They are found in abundance in Guano,

having passed through the alimentary canals of birds which feed

on marine animals. Since they occur in quantity in the surface-
waters of the ocean and of lakes, they constitute a very large
proportion of both the marine and freshwater plankton. Many of
these plankton Diatoms are furnished with long spines or spinous
processes, and they are frequently associated to form free-floating
colonies. Some genera and species are exclusively pelagic in
2
habit. According to Voigt' such forms are sometimes furnished
with gelatinous threads and membranes, by means of which their
floating capacity is materially assisted.
Some of the freshwater species are occasionally the cause of
foulness of drinking water 3 This is due to the formation of an oil
.

1
Castracane in Anrial. de Micrographie, ix, 1897.
'

Voigt in Biol. Centralbl. xxi, 1901.

Whipple & Jackson in Jourii. New Engl. Waterworks Assoc. xiv, 1889.
>
 Bac'dlarieoi 271

analogous to the essential oils, and the foulness could be prevented

by storing water in the dark. It has been suggested that the
immense beds of petroleum which exist in certain regions have
had their origin from the oil secreted in the protoplasm of
Diatoms 1 .

Miquel has made many interesting experiments on the cultiva-

tion of Diatoms, and he finds that, although they thrive in water
at freezing point, they cannot withstand being frozen. Their
vitality is destroyed at 15 C., and a temperature above 45 C. is

rapidly fatal. Desiccation is also fatal. The yellow rays of light

are the most favourable for cultivation, so that cultures should be
placed under yellow glass.
Large numbers of fossil Diatoms are known. Not only are
these minute plants actively engaged at the present time in
forming oceanic and lake deposits, but the numerous Diatumaceous
Earths are a proof of their activity in former ages. These earths
are of a white or grey colour, often so soft and friable as to crumble
readily between the fingers, and they are composed almost entirely
of the siliceous valves of Diatoms. They may have had a marine
or a freshwater origin, and most of the forms contained in the
deposits belong to genera, and many of them to species,
now living.
The deposits have been formed in past times at various periods of
the earth's history, but they appear to be principally associated
with rocks belonging to the Tertiary formations. Some of them
are of economic importance, being used as polishing powders

("Tripoli"), as non-conducting materials, as

absorbents for nitro-
of dynamite (" Kieselguhr "), as a
glycerin in the manufacture
dentifrice, and for other purposes. Although many species usually
occur in deposits of this nature, one is usually predominant, form-
ing the great mass of the material. In some parts of the world,
such as in China, Japan, Siberia, Lapland, and other countries,
certain earthy deposits of a diatomaceous origin are mixed with
meal to make a kind of flour. The best known deposits in the
British Islands are those at Dolgelly in Wales, and at Toome

Bridge in Antrim, Ireland. The deposit at Biln, in Bohemia,

which is about 14 ft. in thickness, was estimated by Ehrenberg to
contain some 40,000,000 of the frustules of Diatoms in every cubic

1
Kramer & Spiller in Berichte Deutsch. Ckern.
Gesellscb. xxxii, 1899.
272 Bacillariece

inch 1
The well-known deposit at Richmond, Virginia, U.S.A., is
.

while on some of
very extensive and reaches a thickness of 30 ft.,
the recent geological surveys beds have been discovered in the
western states of America no less than 300 ft. in thickness. These
earths contain on an average 80 / of Silica.
It is generally appearance of fossil
assumed that the earliest

Diatoms is in the Upper Cretaceous (chalk), although Castracane

has recorded the occurrence of certain species in coal from the
2
stated that he has
English Carboniferous beds, and Edwards has
found valves of Diatoms belonging to the genera Synedra and
Melosira in still older rocks in New Jersey. These observations,
however, require confirmation.
The Bacillariese has been placed by some authors in close
others as an order of the
proximity to the Conjugate and by
of Diatoms are sufficiently dis-
Phseophycea?, but the characters
tinctive and their structure so uniform as to warrant their position
as a distinct class, the affinities of which are doubtful.
I have adopted, with slight alterations, the classification of
Diatoms put forward by Schutt 3 and since followed by Lemmer-
,

mann and others. It is to my mind the most natural one, as it

separates all those Diatoms with a radial symmetry
from those in
which the frustules are zygomorphic or otherwise irregular.
Classifications based upon the disposition and mode of division
4 5
of the chromatophores, such as those suggested by Pfitzer Petit , ,

Pelletan 6 and Ott 7 are impracticable owing to the fact that so

,

many genera and species are unknown in the living state. That
8
published by Prof. H. L. Smith and subsequently adopted by
Van Heurck in his Treatise on the Diatomacea?,' is based upon
'

certain features in the structure of the valves, such as the form of

the raphe, but these characters are not so clear and distinctive

1
In a report on the cheinical composition of the plankton of the Baltic Sea,
Brandt states that 675,000,000 of the dried frustules of Diatoms (mostly Cluctoceros)
weigh one gramme. Cfr Brandt, Beitr. zur Kenntn. der chem. Zusammens. des
'

Planktons,' Wisseusch. Meeresuntersuch., Neue Folge, Bd iii, Heft 2, 1898.

2
Edwards in Amer. Monthly Micr. Journ. xx, 1899, p. 292.
3
Schutt in Engler & Prantl's Die Natiirl. Pnanzenfam. I Teil, i, Abteilung b,
1896.
4
Pfitzer,
'

Untersuchungen liber Bau und Entwicklung der Bacillariaceen,'

Bonn, 1871.
8
Petit in Bull, de la Soc. Bot. de France, Paris, 1877.
6
Pelletan in Journ. de Micrographie, xvi, 1892.
7 Ott in S. B. k. Akad. Wissensch. Wieu, cix, 1900.
8
H. L. Smith in The Lens, Chicago, 1872.
 Badllariece 273

as the primary divisions adopted by Schtitt. The Bacillarieae are

divided into the two following orders :

Order I. Centricce. Valves with a concentric or radiating

symmetry around a central point without a raphe or pseudo-
;

raphe valve-view circular, polygonal, or broadly elliptical,

;

rarely boat-shaped or irregular.

Order II. Pennatw. Valves truly zygomorphic or irregu-
lar, never centric valve-view mostly boat-shaped or needle-
;

shaped, with markings arranged in a sagittal manner in

relation to the raphe or pseudo-raphe (sagittal axis).

Order I. CENTRICLE.
This order includes a relatively small proportion of the known
species of Diatoms, and few even of these are inhabitants of
fresh water.
The cells are commonly cylindrical and when seen in cross-
section (or from the valve-view) they are circular, polygonal,
elliptical, or rarely more elongate. Some forms are disc-shaped,
their diameter being much greater than their length others are ;

ol equal length and breadth or longer than their diameter.

Many
occur as solitary, free-floating individuals, but others are joined
more or less firmly by their valve-facesto form cylindrical filaments
of variable length.
The structure of the valves is typically centric, even in those

species in which the valve-view is zygomorphic in outline. The

markings on the valve-face are either concentrically disposed or

arranged in the form of radiating strise, the latter often termi-

nating in marginal punctulations or dots. In all cases the
arrangement of the markings is in relation to a morphological
centre and never in relation to a middle line. There is no raphe
or pseudo-raphe.
There are usually many small parietal chromatophores of a
but occasionally only two large plates
plate-like form in each cell,
are present.

Auxospores are known only in a few species.

The order is divided into four sub-orders, only two of which include
British freshwater species. One of the sub-orders the Biddulphioideoe
includes a large number of conspicuous marine Diatoms.
Sub-order 1. Discoidece. Cells shortly cylindrical or disc-shaped,
in valve-view circular; hyaline or with radiating or areolated markings.

w. A. 18
274 Butillariete

Sub-order 2. Solenoideoe. Cells elongate, cylindrical or subcylin-

drical, circular or broadly elliptical in cross-section (or in the valve-

view) ; valves exhibiting a scaly structure apices often obliquely

;

conical and furnished with a spine or a hair.

Sub-order 1. DISCOIDE^E.
In this division of the Centricse the cells are more or less

shortly cylindrical and often disc- shaped. In the valve- view or in

cross-section they are circular, and the valve-faces are frequently
very convex. The may cells be solitary and free-floating, or they
may be joined into long chains or filaments by gelatinous cushions.
The valves are generally without any kind of protuberances, and
they may be hyaline or exhibit areolations or radial striations of
any degree of coarseness. Very often the markings on the valves
are divided into distinct sectors, and sometimes there a ring of
is

small spines (rarely of long bristles) at the outer margin of each

valve.
The chromatophores consist of numerous small plates, either
circular in outline or with lobed margins. The nucleus is generally
central, but it may lie near the girdle in a slight thickening of the

primordial utricle.
Only two British families of this sub-order possess freshwater representa-
tives.

Family 1. Melosiracece. Cells subspherical or shortly cylindrical,

circular in cross-section, and united to form long filaments or chains;
girdle usually with a well-marked structure.

Family 2. Coscinodiscacece. Cells mostly disc-shaped and solitary ;

valve-view circular ; girdle usually without structure.

Family 1. MELOSIRACE^.
The cells are mostly shortly cylindrical, rarely subspherical,
and they are usually united by gelatinous cushions to form
filaments of considerable length. The valve-view (and the cross-
section of the cell) is circular or very rarely somewhat compressed.
There is a great uniformity in the type of the cell, and the valves
may be flat, convex, or greatly arched. The valve-face is often
divided into concentric areas, a broad central region and a
peripheral ring of variable width. In some forms there is a
circular keel, occasionally provided with small wart-like protu-
berances, and in others the valves possess a peripheral ring of
 Melo&iracece 275

small spines or teeth. The girdle usually exhibits a well-marked

structure. The chromatophores are small, numerous, parietal and
plate-like.
Genus Melosira Ag., 1824. [Gaillionella Bory ;
Lysigonium
Link; Liparogyra Ehrenb. ;
Orthosira Thwaites; Sphcerophora
Hass.] The frustules are cylindrical, ellipsoidal, or globular, and

Fig. 126. A and B, Melosira arenaria Moore, from Shipley Glen, W. Yorks.
C E, M. varians Ag., from the river Cam at Cambridge; E, showing forma-
tion of auxospore. (All x 450.)

are united to form filaments of variable length. The valve-view is

circular and plainly punctate. Auxospores are formed without
conjugation, by the rejuvenescence of the contents of a mother-cell
to form a much larger daughter-cell, the long axis of which is either

parallel or at right angles to that of the mother-cell. These large

daughter-cells continue to divide while still
remaining parts of the
original filament.

The genus is Sect. 1. Eumelosira Schiitt,

divided into four sections:
including M. arenaria Moore, granulata (Ehrenb.) Ralfs and M. Roeseana
J/.

Rabeuh. Sect. 2. Lysigonium Link, including J/. carians Ag.

;
Sect. 3. ;

Podosira Ehrenb., marine Sect. 4. Gaillionella Bory, including M. numtnu-

;

loides (Bory) Ag.

There are some six or eight freshwater .species occurring in the British
Islands. J/. rarians Ag. is one of the most abundant of the centric Diatoms, oc-

curring in large quantities in ponds, ditches and slow rivers (fig. 126 C E). J/.
arenaria Moore occurs on wet rocks, sometimes forming crisp mat-like masses
on dripping sandstone rocks. It is common on the Carboniferous Sandstone
of England. J/. granulata (Ehrenb.) Ealfs occurs in boggy pools and also in

182
276 BacillariecB

the plankton. M. Roeseana Rabenh. occurs on damp rocks among various

Myxophycese and also among damp mosses. The filaments of
J/. granulata

are sometimes not more than 5p, in diameter, but those of M. arenaria reach
100/i in diameter (fig. 126 A
and B).

Family 2. COSCINODISCACE^.
In this family of the Discoidese the cells are generally disc-
or more rarely highly arched.
shaped, with the valves fiat, convex,
In the valve-view they are circular. There is an absence of warts
or other protuberances, but there is sometimes a peripheral ring of

spines. In some the valves possess concentric areas with different

types of structure, but they are not divided
into sectors by special
radial strands. The usual type of structure consists of radial rows
of punctulations or areolatioiis, and there are never any excentric

spots. The commonly structureless or its structure is

girdle is

almost indeterminable. The cells are solitary and free-floating.

The chromatophores are small and numerous, consisting of rounded
or lobed, parietal plates. Most of the genera of this family are
marine or fossil, and in many instances the distinctions between
them are very obscure. There are only three British genera
inhabiting fresh water.
Genus Cyclotella Kiitz., 1833. The frustules are disc-shaped

and the valves are circular, exhibiting two concentric areas. The
inner area is smooth or granulate, but the outer annular area

radiating stria?, which are smooth

or punctate. Occasion-
possesses
allyminute spines are present near the margin of the disc. The
valves are excentrically bullate in most species, so that the frustule
seen from the girdle-view possesses undulate margins.

There are five species known to occur in the freshwaters of the British

operculata Kiitz. (fig. 127 B and C), G. Meneghiniana Kiitz.,

Islands. C. and
C. Kiitzingiana Chauvin are more or less widely distributed, and C.
comta
(Ehrenb.) Kiitz. is often abundant in the plankton. They vary in diameter
from 10 30/x.

Genus Stephanodiscus Ehrenb., 1845. The valves are

circular, with radiating series of punctulations alternating with

of lines.
radiating smooth spaces which present the appearance
In the centre of the valve there are scattered punctulations and
round the margin is a ring of simple acute spines. The centre of
the valve is
generally bullate.
 Coscinodiscacece 277

St. Hantzschianus Grun. occurs in the plankton of Lough Neagh and in

the river Thames diam. of frustules 12 20/u; fig. 127 A.
;

Fig. A, Stephanodiscus Hantzschianus Grun. (After Schroder, x 544.) B

127.
and C, Cijclotelhi operculata Kiitz., from Shipley Glen, W. Yorks. ( x 450).
D, Coscinodiscus lacustris Grun., from the plankton of Lough Neagh, Ireland
(
x 450).

Genus Coscinodiscus Ehrenb., 1838. The valves are circular,

elliptical, or rarely subrhomboidal, without any striae or costse, but

with radiating punctulations or areolations. There is a distinct

edge to the disc, usually furnished with a ring of submarginal
spines.

The limits of this genus are exceedingly difficult to define and "the
synonymy is most confusing. There are over 300 known species, mostly
marine, and only one occurs in the freshwaters of the British Islands. C.
lacustris Grun. occurs in the still waters of rivers and lakes, and reaches a
diameter of 60/n (fig. 127 D). It sometimes occurs in the plankton of lakes.

Sub-order 2. SOLENOIDE^E.
In this second sub-order of the Centricse the cells are rod-
shaped, many times longer than the diameter, and mostly circular
in cross-section. After division the cells sometimes remain
attached in more or less fragile chains.
278 BacUlariece

Family 1. RHIZOSOLENIACE^E.
The cells are elongated,more or less cylindrical, and often form
chains. In cross-section (or in the valve-view) they are either
circular or broadly elliptical. The extremi-
ties of the cells are attenuated and either
symmetrical or asymmetrical. In the latter
case there is a calyptra or hood surmounted

by a sharp spine or a long needle-like seta.

This attenuation of the extremities of the
cells is really a more or less excentric, coni-
cal projection from the valve-faces. The
girdle exhibits numerous rings of large
rhomboidal scales, and the frustules are
sometimes imperfectly siliceous. The chro-
matophores consist of numerous small plates,
generally somewhat elongated.
Genus Rhizosolenia Ehrenb., 1858 em. ;

Peragallo,1892. The frustules are very

elongated and subcylindrical. The valves
are asymmetrical, terminating in a hood
which is furnished with a spine or a long
seta. The spines or bristles are excentric,
but are disposed parallel to the long axis of
the The girdle region of the cell con-
cell.

sists of rings of scales, which are more or less

imbricate. The chromatophores are small,
rounded or elongated plates. The auxo-
spores are formed without conjugation.
Hypnospores are sometimes produced, one
or two in each mother-cell.

Most of the species of this genus are marine,

Fig. 128. Ehizoso- but two are known to occur in the freshwater
lonijiseta Zach.
(After Schroder, x 544.)
plankton of the British Islands. R. longiseta Zach.
A, frustule showing rest- (fig. 128)
occurs in the plankton of Loch Shin and
ing spore; B, germina- Loch a Gharbh Bhaid Mhoir, Sutherland, and
tion of resting spore.
R. eriensis H. Smith also occurs in Loch Shin.

Genus Cylindrotheca Rabenh., 1859. The frustules are

symmetrical and spindle-shaped, with the apices much attenuated.
 Rhizosoleniacece 279

They are ornamented with several running along the

spiral lines,
walls of the valves from end to end of the frustuleand crossing one
another at intervals. The chromatophores are small and granular.
The structure of the cell-wall is only imperfectly known and the
position of the genus is uncertain. It should perhaps be associated
with the Nitzschiacese.
C. ff
radii's (Breb.) Grim. = Nitzschia Tcenia W. Sin.] is the only known
\_

species and is a rare Diatom in the fresh and brackish waters of England.

Order II. PENNAT^E.

This is by far the largest order of the Bacillariese and includes
allthose Diatoms in which the valves are not of a centric type.
The structure of the valves is arranged in relation to a line and
not to a central point. The cells are acyclic, being rod-shaped,
elliptic, or
boat-shaped in cross-section
(or valve-view). The
marks (striae or costse) on the valves are disposed on either side of
a sagittal line (which is either a raphe or a pseudo-raphe), some-
times arranged at right angles to it and sometimes forming an
acute angle with it.
The cells exhibit great variability of form and in most cases are
truly zygomorphic. The commonest type of cell is the naviculoid
or boat-shaped type, but the frustule may be a flattened plate or
an elongated rod, which may be straight, arcuate, or sigmoid.
The different forms exhibit all degrees of development of the
raphe. In the lowest forms of the Fragilarioidere it is entirely
wanting, but in other forms of the same group there are the first
beginnings of a pseudo-raphe. In the Naviculoideas the raphe is
present on both valves of the frustule, and it is sometimes situated
on a median or obliquely disposed keel. It reaches its highest
development in the NaviculaceaB.
There is a striking absence of spines and long processes from
the frustules of Diatoms of this order, although in a few instances
they are furnished with small prickle-like excrescences.
Great variation is exhibited in the size and form of the chro-
matophores, which are in all cases parietal. In the lower families
of the Pennatas each cell contains a large number of small plates
(the <coccochromatic arrangement), but in the higher families
one or few large lobed plates occupy the greater part of the inner
surface of the cell -wall (the placochromatic arrangement).
280 Bacillariece

The methods of formation of auxospores are as yet only

imperfectly known. In some of the higher forms the method
is a sexual one, and the
highest known type is found in the
Surirellacese.

The order is divided into the following sub-orders :

Sub-order 1.
Fragilarioideoe. Cells mostly straight, rod-shaped or
lanceolate, without a raphe, but sometimes with a pseudo-raphe or
showing indications of the commencement of a raphe.
Sub-order 2. Achnanthoidefe. Cells crooked or suddenly bent,
with a raphe on one valve and a pseudo-raphe on the other.
Sub-order 3. Naviculoidece. Each valve of the cell with a raphe ;

valves without a keel (or rarely with a keel) in the sagittal line (line of
raphe).
Sub-order 4. Nitzschioideoe. Each valve of the cell with a raphe,
which is situated in a sagittal keel with carinal dots. The keels of the
two valves are situated on opposite sides of the frustule or displaced
to the same side. Cells in transverse section rhombic.

Sub-order 5. Surirelloidece. Each valve of the cell with a pseudo-

raphe and generally with spreading submarginal wings valves strongly
;

costate.

Suborder 1. FRAGILARIOIDE^.
In this sub-order the cells aremostly rectangular in the girdle-
view, and rod-shaped, lanceolate, or arcuate in the valve-view. In
the Meridionacese they are elongate and cuneate in both views.
Sometimes they occur as solitary individuals, but they are more
commonly joined by gelatinous cushions into either straight or
zig-zag, ribbon-likefilaments. In some there are false septa
traversing the frustules, but in others these are absent. There is
no raphe, but in most of them there is a pseudo-raphe forming a
sagittal line in relation to which the markings of the valves are
arranged. In the Meridionaceas the presence of this pseudo-raphe
is
scarcely evident. The chromatophores are mostly numerous,
small and granular, but in the Eunotiacea3 they are few in number,

large and plate-like.

The sub-order includes the following five families :

Family 1. forming tabular plates in the

Tabellariacece. Cells

girdle-view, united to form band-like or zig-zag filaments; with well-

developed false internal septa. Valves with a straight median pseudo-
raphe.
 Tdbellariacece 281

Family 2. Meridionacece. Cells elongate and cuneate, united to

spiral filament or situated on branched gelatinous

form a flat stalks ;

with false septa. Valves with a very indistinct median pseudo-raphe.

Family 3. Diatomuceie. Cells mostly rod-shaped or subrectan-

gular, united into band-like or zig-zag filaments; valves with feebly

developed transverse septa and a median pseudo-raphe.
Family 4. Fragilariacew. Cells in girdle-view rectangular, gene-

rally united to form ribbon-like filaments. Valves without any false

septa and with a median pseudo-raphe.
Family 5. Eunotiacece. Cells slightly curved or arcuate in the

valve-view, solitary or united into band-like filaments. Pseudo-raphe

nearer to one edsre of the valve.

Family 1. TABELLARIACE.E.
The cells are stout and expanded in the girdle-view to form

flat, rectangular plates.In the valve-view they are bilaterally

symmetrical, linear or linear-elliptic, often with a swollen median
portion and sometimes with subcapitate extremities. They are
mostly united to form band-like or zig-zag filaments by mucous
cushions on the valve-faces or at the angles. In the interior of
the frustules are two or more false longitudinal septa. Both valves
are precisely similar with a straight median pseudo-raphe (sagittal
line). The chromatophores are numerous and granular.
Genus Tetracyclus Ralfs, 1843 em. Grun., 1862.
;
The
frustules are tabularand are united to form short or long, ribbon -
like filaments. There are a number of perforated longitudinal
septa, which appear in the girdle-view as ribs
with thickened
apices. There are also false transverse septa at right angles to
these, which appear as few or many costse (or ribs) in the valve-
view. The median portion of the valve may or may not be
swollen.

There are two British species, T. lacustris Ralfs (length of valves up to

30 p.;fig. 129
A C), which prefers hilly districts and is often found in the
plankton of mountain lakes, and T. rtipestris (A. Br.) Grun., a species which
occurs on dripping rocks in mountainous areas.

Genus Tabellaria Ehrenb., 1839. The frustules are tabular

and united to form zig-zag filaments, the basal cell often being
fixed to the substratum by a mucous cushion at one corner of
the valve. There are a number of perforated longitudinal septa,
which appear in the girdle-view as prominent lines which fail to
282 Bacillariece

reach the middle of the cell. Sometimes instead of a large

perforation in a septum, the septa are alternate, and only extend
about two-thirds the distance from one end of the valve to the
centre. The valve-view is sublinear, more or less strongly swollen
in the middle and subcapitate at the extremities; its .surface is

transversely striated, the striations being slightly broken in the

middle, thus indicating the presence of a rudimentary pseudo-
raphe. The auxospores are formed two in each mother-cell.

Fig. 129. A C, Tetracyclus lacustris Ralfs, from the plankton of Loch Shin,
Sutherland. D and E, Tabellaritt fenestrata (Lyngb.) Kiitz., from Mickle Fell,
N. Yorks. F and G, T. flocculosd (Roth) Kiitz., from Mickle Fell, N. Yorks.
(All x500.)

There are two common British species, of which T

flocculosa (Roth) Kiitz.
.

(tig. 129 F and G) is the most abundant. T fenestrata (Lyngb.) Kiitz. (fig.
129 D and E) has more slender and elongate valves, the largest specimens
noticed having reached a length of 137 /u.
1
. A pretty variety of the latter
species var. asterionelloides which the frustules are arranged in
Grun. in
circles, is known from the plankton of Loughs Neagh and Beg in Ireland, and
from that of many lochs in Scotland and in the English Lake District. It
is
probably general in the British freshwater plankton and there are two very
distinct forms of it.

Genus Diatomella Grev., 1855. The frustules are tabular,

solitary, or joined to form ribbon-like filaments. There are only
1
West & G. S. West in Trans. Roy. Irish Acad. 1902, vol. xxxii, sect. B, part 1,
p. 90, t. iii, f. 8.
 Meridionacece 283

two false longitudinal

septa, well seen in the rectangular girdle-
view, and which are perforated by three openings, one central and
two polar. The valve-view is oblong-lanceolate, with a slight
median swelling its surface is transversely striated with indications
;

of a pseudo-raphe and slight traces of nodules.

D. Balfouriana Grev. is a rare British Diatom the frustules of which

reach a length of 30 p.

Genus Denticula Klitz., 1844. The frustules are more or less

tabular, solitary or attached to form short filaments. There are a
row of imperfect transverse septa in each valve, which appear as
capitate marginal ribs in the subrectangular girdle-view. The
valve- view is lanceolate and the septa appear as strong transverse
ribs,between which are delicate punctate striae. The face of each
valve is carinate and for this reason Denticula has been associated

with Nitzschia. In the region of the girdle, and between it and

the valve on either side, is a longitudinal septum with a row of
perforations, the edges of these perforations being fused to the
transverse septa. There is no raphe or pseudo-raphe.
D. elegans KiAtz. and D. tennis Kiitz. (tig. 130 C and D) are both frequent
amongst mosses on wet or dripping rocks. Valves up to 45 ^t in length.

Family 2. MERIDIONACE.E.
The frustules are more or less rod-shaped and cuneiform, and
by their valve-faces to form a flat spiral filament,
are either united
which often makes two complete turns, or they are disposed at the
extremities of a branched system of gelatinous stalks. In both
the valve- and girdle-views the frustules are symmetrical in
relation to a longitudinal axis, but asymmetrical about a transverse
axis. In some there are two (sometimes more) false longitudinal
septa, but no transverse septa; sometimes these are confined to
the broad end of the frustule (I/icmophora) and at other times
they extend the whole length of the valves (Climacosphenia). In
others there are no false longitudinal septa, but numerous short
transverse septa pass across the keeled face of each valve (Meridian).
There is no raphe, but there is a structureless sagittal line which
indicates the firstcommencement of a pseudo-raphe. Reaching
almost up to this clear line are numerous transverse striations of
a very delicate character. The chromatophores are scattered in
284 BadllariecB

the form of numerous small plates. Two auxospores arise from

two mother-cells.
Genus Meridion Ag., 1824. The frustules are very similar to
those of the genus Diatoma. The valve-view is clavate, sometimes
with a subcapitate apex, and the girdle-view is cuneate with a

Fig. 130. A and B, Meridion circular e (Grev.) Ag., from Shipley Glen, W. Yorks.
(x500). C and D, Denticula tennis Kiitz., from Wicken Fen, Cambridge
(
x 600).

truncate base and apex. There are no false longitudinal septa,

but a number of imperfect transverse septa, which appear as very
short marginal costse in the girdle-view and as transverse costce in
the valve-view. Between these costa? are fine punctate striae
interrupted in the middle by a smooth line or pseudo-raphe. The
frustules remain attached after division, forming beautiful, fiat,

spiral filaments, which are free-floating.

M. circulare Ag. (fig. 130 A and B) is often abundant in stagnant ditches
and ponds, and may frequently be obtained in pure masses. The var. con-
strii-tum (Ralfs) V. H. is also frequent. It is a Diatom which appears to be
most abundant in the early spring, often forming brown flocculeut masses
around submerged grass-leaves, etc. It is one of the prettiest of British
freshwater Diatoms length of valves up to 25 p.
;

Family 3. DIATOMACE^.
The frustules are elongate, rod-shaped or lanceolate in the
valve-view, and are united to form ribbon-like or zig-zag filaments.
There is no keel, but the valves possess strongly marked transverse
ribs which project inwards as more or less deep transverse septa.
There are no false longitudinal septa. The pseudo-raphe is con-
spicuous or it may be wanting. The girdle-view is rectangular.
Genus Diatoma D.C., 1805. The valve- view is lanceolate or
linear, sometimes with capitate extremities, and is furnished with
Fragilariacece 285

E
280 JBaoillariece

fine or rather coarse and composed of bead-like punctulations.

The girdle-view is
rectangular. The frustules are joined together
by their valve-faces to form ribbon-like filaments, or they are
united by small mucous cushions at their corners to form zig-zag
filaments. In the section Eufragilaria Ralfs the pseudo-raphe is
very insignificant and the chromatophores are small and granular ;

in the section Staurosira Ehrenb. (= Odontidium Kiitz.) the

is broad, often lanceolate, and the chromatophores
pseudo-raphe
are plate-like as in Synedra.

F. capucina Desmaz. (length of valves 30 60 p. tig. 132 C and D) is

;

much the commonest freshwater species, but F. mutabilis (W. Sm.) Grun.
and F. virescens Ralfs are also general. F. comtruens (Ehrenb.) Grun. and
F. Crotonensis (A. M. Edw.) Kitton are rarer freshwater species.

Genus Synedra Ehrenb., 1831. The frustules are much

elongated and occasionally bent or somewhat undulated. The
valve- view is
commonly linear or
linear-lanceolate, with obtuse or

subcapitate extremities, and there

is
generally a pseudo-raphe or a
hyaline sagittal line. In the centre
of the valve-face there is generally
a small rounded hyaline space, and
sometimes central and polar nodules
are present. The valves are finely
striated, the stria; being transverse
and reaching up to the pseudo-
raphe on each side. The girdle-
view is elongated with truncate
apices. Each frustule contains two
plate-like chromatophores with un-
dulated or indented edges. Most
of the species are but
A solitary,
some occur clustered in radiating
Fig. 132. A and B,
pulchella Kiitz.,from Cambridge or fan-shaped colonies.
(x500). C and D, Frtiiiilnria
rhcre are about ei g ht British fresh-
<-<ipii,-ina Desmaz., from Shipley,
W. Yorks. (
x 520). water species, several of which are
common, being found in almost every
kind of suitable locality and often occurring in immense abundance in the
waters of lakes and springs. S. Ulna (Nitzsch) Ehrenb. (length of valves
fig. 132 A and B) and
150 250 /x), <S'. pulchella Kiitz. (length 60 ISO/* ;
 Eunotiacece 287
.y. Acus (Kiitz.) Grun. are the most almndant. variety of S. Ulna A
var. splendens (Kiitz.) V. H. is a very striking object, the valves Ix-in.^

arranged in radiating groups and reaching a length of 340 p.. <S'. capitata
Ehrenb. is general, but not so common as the previous species.

Genus Asterionella Hassall, 1850. The frustules are narrow

and linear, with swollen
apices both in the valve-
and girdle-views they ;

are grouped in radiating

coloniesinone plane, like
the spokes of a wheel,
their basal extremities

being attached by deli-

cate mucous cushions.
In the valve-view the
cells are narrowly linear-
fusiform with capitate
apices, one apex being-
larger than the other.
In the girdle-view the
cells are linear with
Fig. 133. Asterionella formosa Hass., from
swollen, truncate ex-
the plankton of Lough Neagh, Ireland ( x 450).
tremities. The valves

possess delicate transverse striations,

and there is a median pseudo-
raphe and a hyaline area in each capitate apex.
A. formosa Hass. (fig. 133) is a common species in the quiet waters of
ditches, ponds and lakes; length of valves 65 90 p.. A. granllima Heib. is
narrower and more elongate (length of valves up to 130 /*), and with the
preceding species, of which it is perhaps only a variety, is a regular and
abundant constituent of the British freshwater plankton. The colonies of
this genus are somewhat fragile and they easily become dismembered.

Family 5. EUNOTIACE^.
The cells are free-floating and either solitary or united by
their valve-faces to form ribbon-like filaments. The frustules are
more or less curved or arcuate in the valve-view, the dorsal border
of which is often undulate, the ventral margin being concave and
with or without a central swelling. There is either a reduced
raphe or a pseudo-raphe close to the ventral or concave margin,
and the valve is transversely striated. The central nodule is
288 Bacfllariew

nodules are disposed near the ventral

mostly absent, but the polar
The girdle-view is generally rectangular. Each frustule
margin.
possesses two, parietal, plate-like chromatophores.
Genus Ceratoneis Ehrenb., 1840. The cells are solitary and
in the valve-view the ventral or concave side exhibits a more or
less prominent central swelling.
The apices of the valves are
obtuse, subcapitate, or rostrate-capitate. There is a well-marked
the ventral margin and interrupting the
pseudo-raphe close to
transverse strise. The polar and central nodules are comparatively
distinct. The girdle-view is linear-rectangular.

C. Arcus (Ehrenb.) Kiitz. [ =

Eitnf<' A ><><* W. Sin.], with obtuse apices, is
a frequent Diatom in mountainous areas, particularly in mountain streams ;

length of valves 35 134 D.

60/*; fig.
A
variety of it var. Amphioxys
(Rabenh.) De Toni distinguished by produced and subcapitate apices, is
its

also abundant in mountainous and pure gatherings of it are by no

districts,
means uncommon. The valves reach a length of about 85 p..

Genus Eunotia Ehrenb., 1837. [Himantidium of various

authors ;Desmagonmm Ehrenb.,
1848 ; Amphicampa Ehrenb.,
1849 ;
Pseudeunotia Grim.,
1865 ;
Climacidinm Ehrenb.,
1867.] The valve-view is ar-

cuate or bow-shaped and the

dorsal margin is often undulate
or nodulose. The apices are
obtuse or more often subcapi-
tate. There is no central nodule
and the pseudo-raphe is not
very evident owing to its posi-
H '4
tion along the ventral margin.
B W| The transverse stria? are in-
terrupted across the valve-face.
Fig. 134. A, Eunotia pectinalis (Kiitz.) The girdle-view is rectangular.
Eabenh. var. itndulatn Ralfs, from Bail-
don, W. Yorks. ( x 600). B, E. robust a
In one section of the genus
Ealfs, from Dolfielly, Wales ( x 600). C,
{Himantidium) the cells are
E. jirnciliti (Ehrenb.) Rabenh., from
Lenvick, Shetlands (x500). D, Cera- united by their valve-faces to
toneis Arcus Kiitz., from Cautley Spout, form long, flexuose, ribbon-like
W. Yorks. (
x 520).
filaments. In the other section

(Eunotia) the cells are generally solitary, but they may be epiphytic
in clusters on other larger Alga 1
.
 Achnanthacece 289

In the E. gmcilis (Ehrenb.) Rabenh. (fig. 134 C) and E. pec-

first section,

Rabenh. are the two commonest species. In the second section,

(Kiitz.)
E. lunaris (Ehrenb.) Gran, is much the commonest species (length of valves
50 90 p.), but E. tetraodon Ehrenb., E. Veneris Kiitz. and E. biceps nob.
[
= Si/,,<'(lr<t biceps W. SIM.; E. flexuosa var. bicapitata Grun.] are general
in
mountainous areas.

Sub-order 2. ACHNANTHOIDE^.
The Diatoms of this sub-order are at once distinguished by the
crooked or geniculate character of the frustules. Seen in the
girdle- view each frustule is suddenly bent in its median portion,
a character not exhibited by any other group of Diatoms. Another
important feature of the group is the presence of a raphe on one
valve of the frustule and a pseudo-raphe on the other. Only two
families are included in the sub-order.

Family 1. Achnanthacece. Frustules elongate and more or less

rod-shaped, generally occurring as stalked epiphytes.

Family 2. Cocconeidacece. Frustules flat and plate-like, subcir-

cular or broadly elliptical in outline, occurring as epiphytes attached
by their flat valve-faces.

Family 1. ACHNANTHACE^I.
The and symmetrical in outline in the
frustules are straight
valve-view, but geniculate and asymmetrical in the girdle-view.
In the valve-view they are in all cases linear or fusiform, often
with capitate apices. One valve possesses a pseudo-raphe without
any trace of nodules, whereas the other valve possesses a true
raphe with both central and terminal nodules. The valves are

transversely striated, the consisting of fine rows of punctse,

striae

and in some species costse are present between the rows of punctae.
The chromatophore is usually a thick parietal plate, mostly covering
the convex valve, but in some species there are a number of small

granular chromatophores.
Genus Achnanthes Bory, 1822. [Achnanthidium Klitz., 1844 :

Grim., 1880.] The Diatoms of this genus possess the essential

features of the family, and are either free-floating and solitary or
stalked and There are two sub-genera, Euaclimnithes
epiphytic.
w. A. 19
290 Batillariece

Schiitt, in which the raphe and pseudo-raphe are straight and

median or a little excentric; and Acli-
nttnthidium Klitz., in which the raphe
and pseudo-raphe are sigmoid. In some
species there is a distinct stauros on
the lower valve. Auxospores may be
formed in two ways two from a mother-
:

cell without
conjugation, or one formed
by the conjugation of two cells.
The largest freshwater species with a straight
raphe, and also rather a rare one, is ^4. coarctata
Breb. ; length of valves 18 43 p. A. exilis
Kiitz., A. linearis W. Sm. and A. microcephala
F E Kiitz. are very small species which are often

Fiff. 135. AAchmin-

C, abundant, frequently occurring as stalked epi-
thes Hungarian Grnn., from
phytes in thick masses round other filamentous
near Sutton, Cambridge x A1 8e> A flexella (Kiitz.) Breb. possesses a
520). D F, Cocconeis Pla-
(

... ,
, ... .

and often occurs in abundance

J

centula Ehrenb., from Sheep's figmoid raphe

Green, Cambridge ( x 520). in hilly districts length of valves 35
;
50 //.

Family 2. COCCONEIDACE^.
The frustules are symmetrical in both the valve-view and in
transverse section, as in the family Achnanthaceas. They are,
however, flat and plate-like, exhibiting a broadly elliptical or

subcircular outline in the valve-view. The valves are dissimilar,

one possessing a pseudo-raphe and the other a raphe with a central
nodule, but polar nodules are generally absent. The frustules
frequently possess perforated transverse septa which appear in the
valve- view as ribs. The girdle- view is generally more or less bent
or arched. The cells possess one chromatophore, which is
mostly
parietal on the convex valve.
Genus Cocconeis Ehrenb., 1835. [Orthoneis Grim., 1868 ;

Anorthoneis Grun., 1868.] This is the only genus of the family

containing any freshwater species. The frustules are flat, plate-
like or leaf-like, and somewhat the shape of a concavo-convex lens ;

they possess punctate striae, transverse in the middle of the valve

and radiating towards the poles. The costre seen in the valve-view
of some species and which represent the false septa, are confined
to the outer border of the valve. One auxospore is formed from a
single mother-cell.
 Naviculacece 291

There are many marine species of this genus, but only two C. Pediculus
Ehrenb. and C. Placentula Ehrenb. (fig. 135 D F) inhabit fresh water, both
of them being abundant and widely distributed all over the British Islands.
C. Pediculus is distinguished from C. Placentula by the form of the valves,
which are more attenuated towards the poles, and by the different markings.
They are both of approximately the same size (12 35 p. in length) and occur
as epiphytes on filamentous Algse and other water-plants. Filaments of
Cladopkora and Vaucheria are often completely covered with these epiphytes,
which are attached by their flat valve-faces.

Sub-order 3. NAVICULOIDE^E.
This is much the largest sub-order of the Pennatse and is

characterized by the presence of a true raphe on each valve of the

which is furnished with a central and two
frustule, polar nodules.
The valves are usually without a keel, but if present it is situated
in the line of the raphe (sagittal line). The frustules may be
free-floating, occurring large in numbers in mucous tubes, or
attached to some fixed object by hyaline stalks. There are three
well-marked families.

Family 1. Namciilacece. Valves straight, rarely oblique or sig-

moid, generally with a perfectly straight raphe, more rarely with an
oblique one mostly symmetrical about a longitudinal axis (line of
;

raphe) and a transverse axis; girdle-view generally symmetrical and

rectangular.

Family 2. Gomphonemacece. Valves mostly straight (rarely curved)

and symmetrical about a longitudinal axis (line of raphe) in both ;

valve- and girdle-view wedge-shaped.

Family 3. Cocconemacece. Valves always curved, symmetrical

about one longitudinal plane and a median transverse plane valve- ;

view generally curved, often sublimate girdle-view straight and sub-

;

rectangular.

Family 1. NAVICULACE^E.
The Diatoms of this family are exceedingly numerous and
well-defined. The frustules are generally straight and symmetrical
in three planes at right angles to each other (Navicula, Vanheurckia,

etc.), or they may be sigmoid (Gyrosigma), or twisted (Rhoicosigma).

The valves are almost always elongated, although in a few genera

(Amphiprora} they may be relatively short. The raphe is somewhat

variable it is generally median with central and polar nodules,
;

but it may take up a curved position according to the form of the

192
292 Baeittariece

valve. In some the central nodule forms a wide stauros (Stauro-

neis) and in others it is enclosed between two siliceous ridges

(Vanheurckia). In a few genera the valves are carinate, or they

are unequal, with a straight margin and an inflated one. The
valves are striated, the striae being commonly transverse, punctate,
and interrupted by the raphe. In Mastogloia the valves are
of an ordinary striated valve superposed
compound, being composed
on a perforated plate. The frustules are mostly solitary and free-
floating, but in some
forms they are enclosed within a tough mucous
which is often much branched others are attached to a
envelope ;

substratum by hyaline stalks. The chromatophores generally

consist of two large parietal plates which are principally disposed
over the walls of the girdle-faces. Two auxospores are commonly
produced by the conjugation of two mother-cells, which surround
themselves with a wide mucous investment.

There are seven British freshwater genera of the family, distinguished as

follows :

A. Valves simple.
* Frustules
straight and symmetrical in three planes at right angles ;

raphe straight.
t Eaphe straight and simple, with polar and central nodules.
Central nodule small Navicula.
|

\\ Central nodule forming a stauros

Stauroneis.
tt Raphe straight, enclosed between two longitudinal ribs.
Central nodule small Vanheurckia.
I

\l Central nodule linear and greatly elon-

gated Amphipleura.
** Frustules sigmoid raphe sigmoid
; Gyrosigma.
*** Frustules twisted; with sigmoid keel in the

sagittal line; raphe sigmoid Amphiprora.

B. Valves composed of two superposed plates Mastogloia.

Genus Navicula Bory, 1822. [Pinnularia Ehrenb., 1843;

Schizonema Ag., 1824 Colletouema Breb., 1849 Diadesmis Kiitz.,
; ;

The frustules are solitary and free-floating

or enclosed in
1844.]
mucous tubes, rarely united by their flat valve-faces to form ribbon-
like filaments. The valves are quite straight and symmetrical
with regard to the line of the raphe (sagittal line). The raphe is
In the girdle-view the
straight with central and polar
nodules.
frustules are straight and subrectangular. In the valve-view the
form of the very variable, although the predominant shape
cell is

is lanceolate or fusiform. The markings on the valves are more or

 Naviculacece 293

less transverse, sometimes somewhat radiating, and are arranged

in relation to the line of the raphe. They consist of striae or
costae of variable strength and are frequently composed of rows of
punctae. The never quite reach up to the raphe, and some-
striae

times there a hyaline area of considerable size on each side of

is

the central nodule. In rare cases the striae are interrupted by a

smooth longitudinal area on each side of the raphe and parallel to
it. The chromatophores consist of two large parietal plates and in
the formation of auxospores two spores are formed by the con-
jugation of two mother-cells.
Navicula is much the largest genus of Diatoms, or indeed, of
any group of Algae, embracing upwards of 1,000 species, which

Fig. 136. A, Navicula alpina (W. Sm.) Ralfs, from Lerwick, Shetlands (x400).
B, X. iiridis Kiitz., from Baildon, W. Yorks. (x400). C, JV. sphcerophora
Kiitz.,from Wimpole Park, Cambridge ( x 500). D, N. serians (Breb.) Kiitz.,

from Mickle Fell, N. Yorks. ( x 400). E, Stauroneis Pluxnicenteron (Nitzsch)

Ehreub., from Adel Bog, W. Yorks. ( x 400). F, St. acuta W. Sm., from
Shipley Glen, W. Yorks. (
x 450).

occur widely distributed in fresh, brackish and salt water. Many

are also known in a fossil condition. It has been subdivided by
Cleve into the three subgenera Navicula, Stauroneis and Dictyoneis.
The first of these subgenera is divided into the two groups
Eunavicula and Schizonema, and there are twenty-two sections of
the first group, twelve of which contain British freshwater species ;

in the second group there are two sections, one of which contains
294 Banllariece

freshwater species the second subgenus Stauroneis I prefer to

;

regard as a genus and in the third subgenus there are three

;

sections which are exclusively marine.

There are about 70 freshwater species occurring in the British Islands,

principally in stagnant waters. The most striking are N. cuspidaia Kiitz.,
N. nobilis Ehrenb., N. major Kiitz., N. alpina Ralfs (fig. 136 A) and N. lata
in elevated
Breb., the two last-mentioned species preferring boggy tracts
regions, in which localities they are sometimes frequent. N. perpusilla
Grun.
7
(length 12-5 p.) and A gallica (W. Sm.) V. H. (length
,
p) are amongst 815
the smallest British species N. nobilis Ehrenb. is the largest species (length
;

200 400^,) and N. viridis Kiitz. (tig. 136 B) is perhaps the commonest.
Some of the small and abundant species, such as N. exilis Grun. and others,
are remarkable for the rapidity of their movements. Some of the species of
this genus are utilized as test-objects for the objectives of microscopes.

Genus Stauroneis Ehrenb., 1843. [Pleurostauron Rabenh.,

1859 Schizostauron Grun., 1867.]
;
This genus is at once dis-
tinguished from Navicula by the form of the central nodule, which
is transversely dilated to form a stauros. The frustules may be
solitary and free-floating or attached by their valve-faces to form
short filaments.

There are about six freshwater species occurring in Britain, of which /SY.

Phcenicenteron Ehrenb. is the largest and most frequent ;

length of valves
100170^; fig. 136 E.

Genus Vanheurckia Breb., 1868. [Frustulia as amended by

Rabenh., 1851, but scarcely the Frustulia of Agardh, 1824.] The
frustules, which are free-floating or occasionally arranged in a
linear series in a mucous same type
tube, are of precisely the
as those of Navicula. They possess, however, a very distinctive
feature in the sagittal line of the valves. There are distinct polar
and central nodules, which are elongated and enclosed along with
the raphe between two parallel siliceous ribs. The valves are
transversely striated, the stride being very fine and parallel, rarely
slightly radiating in the median portion of the valve.

There are only two British species, V. rhomboides (Ehrenb.) Breb.

[= Navicula rhomboides Ehrenb.; fig. 137 A and B] and V. mdgaris (Thw.)
V. H. V. rhomboides var. Saxonica (Rabenh.) G. S. West = Navicula \_

crassinervia Breb.] is very abundant and widely distributed in the British

Islands, being one of the most frequent Algse in boggy districts; length of
valves 50 80 /i.
Pure gatherings of it can often be collected from Sphag-
num-pools.
 295

Fig. 137. A and B, Vanlieurckia rltoinhoides (Ehrenb.) Breb., from Mickle Fell,
N. Yorks. (x520). C, Amphipleura pellncida, Kiitz., from Chippenharn Fen,
Cambridge ( x 520). D and E, girdle- arid valve-views of Naviculu viridix Kiitz.,
to show chromatophores (ch) and nucleus (n), x 400. F, Eunotia gracilis
(Ehrenb.) Babenh., girdle view to show chromatophores and nucleus ( x 400).

Genus Amphipleura The frustules are solitary

Kiitz., 1844.
and free-floating, elongate-fusiform in shape, with a small marginal
keel near each edge of the valves. The valve-view is narrowly
lanceolate and the central nodule is greatly elongated, forming a
narrow rib which separates the raphe into two short portions, each
portion being situated towards one extremity of the valve and
enclosed between two parallel ribs. These two ribs unite together
at each extremity of the valve to form the polar nodules. The
valves are transversely striated and in some species these strige
are exceedingly fine.

Only one species, A. pelfm-iili Kiitz., inhabits the freshwaters of the

British Isles. It is generally distributedand often abundant length of ;

valves 80 140 137 C. The strife of this species are extremely fine
/A; fig.

(about 37 in 10 /*) and the valves are used as microscopic test-objects. Some
very large forms of it sometimes occur in the freshwater plankton.

Genus Gyrosigma Hassall, 1845. [Pleurosigma W. Sm., 1853;

?
Scalprum Corda, 1835.] The frustules are elongated, of the
naviculoid type, and the valves are convex. In the valve-view
they are sigmoid with obtuse or attenuated extremities. The
There are two sets of striations on the
'

raphe also sigmoid.

is

valves, which cross one another either at right angles or obliquely.

The girdle-view is generally straight and linear-oblong in shape.
The chromatophores consist of a pair of large jagged or perforated
296 Baeillariece

plates disposed in the manner common to all the genera of the

family Naviculacese.
There are only four British freshwater species, of which G. attenuatum
(Kiitz.)Eabenh. (length of valves 190 250 /z; tig. 138 A) and G.
Spencerii
(Queck.) O. K. (length of valves 80 130 /x)
are the most abundant.

Genus Amphiprora Ehrenb., 1843 em. Cleve, 1891. [Amphi-

;

campa Rabenh., 1864.] The frustules

are free-floating and
solitary, with a
around the longitudinal axis.
slight twist
In the valve-view they are of a navi-
culoid form with a sigmoid
raphe, the
latterbeing situated on a prominent
sigmoid keel in the sagittal axis. In
the girdle-view the frustules are
broadly
inflated, with a median constriction and
truncate ends. The girdle itself exhibits
a slight twist, and seen
obliquely the
sigmoid keels of the valves cause the
frustules to appear much more twisted
than is truly the case. The terminal no-
dules are not very conspicuous and there
is a small central nodule. The valves are
finely and transversely striated.

A. paludosa W. Sin. is often found in

fresh water, sometimes in great abundance in
small ponds and ditches length of valves;

Fig. 138. A, Gyrosigma 40 138 B and A. ornata Bailey

C.
atteniifitum (Kiitz.) Rabeuh.,
80/*; fig.

from Chippenham Fen, Cam- is also known from the freshwaters of England.
bridge (x400). B and C,
Amphiprorapaludosa'W.SiQ.., Genus Mastogloia Thwaites, 1848.
from Wimbledon Common,
Surrey (
x 400). The frustules are of a iiaviculoid form
and are enclosed in a gelatinous envelope
of considerable size. The valve-view
is usually elliptic-lanceolate, generally
with produced extremities, and the
girdle-view is
sub-rectangular. Each
frustule possesses two longitudinal septa
with a large central perforation and a
Fig. 139. Mastoyloia row of marginal ones on each side.
Smithii Th\v., from Baildon,
W. Yorks. (
x 500).
These two longitudinal septa are per-
 2i)T

forated siliceous plates situated between the girdle and each valve.
The valves are transversely striated, the striae radiating somewhat
in the centre, and there is a straight median raphe with central
and polar nodules.
About four species are known from the freshwaters of the British Islands,
of which JA Smith ii Thwaites (length of valves 30 45 /i fig. 139) and .I/.
;

Dansei Thwaites are the most generally distributed.

Family 2. GOMPHONEMACE^J.
This family of the Naviculoidese is characterized by the wedge-
shaped form of the frustules, especially when seen in the girdle-view.
In the valve-view they are naviculoid in form, with one pole
generally much larger than the other and with
sinuate margins.
The valves are symmetrical about the sagittal axis (line of raphe)
and the is straight
raphe and median. Longitudinal septa are
present between the valves and the girdle, but they
do not extend
far into the cavity of the cell. Each frustule possesses a large
which is somewhat sinuate and fits closely
parietal chromatophore,
along one girdle-face, covering the valve-faces and most of the
other girdle-face. The frustules are generally attached by their
inferior (or smaller) extremities to a branched system of hyaline

stalks, which is attached to a sub-

stratum. There are two
only
genera, Gomphonema, inwhich the
frustules are straight, and Rhoico-

sphenia, in which they are curved.

Two auxospores are formed from
two mother-cells without conjuga-
tion and grow parallel to each other.
Genus Gomphonema Ag.,1824.
\Gomphonella Rabenh.; Gomphoneis B
Cleve.] The frustules are of vari-
Fig. 140. AandB, Gomphonema
able form in the valve-view, gene- fjemiiiatum (Lyngb.) Ag., from Caut-
'ley Spout, W. Yorks. (
with one extremity conspicu- x 400). C,
rally G. cunstrii-tum Ehrenb., from Chip-
ously larger than the other, often x
peuham Fen, Cambridge ( 400).
fusiform in outline or sinuate at
the margins. Each valve possesses a straight median raphe, and
the central and polar nodules are well-marked. The valves are
generally strongly striated, the strise more or less radiating from
298 BadllariecB

the sagittal axis and in some forms consisting of distinct beads.

In the girdle-view the frustules are cuneate in outward form.
There are about 14 British species of the genus, some of which, such as
G. constrictum Ehrenb. (fig.
140 C), G. acuminatum Ehrenb., G. parvi/l n m
Kiitz. others, are common in every part of the British Islands, frequently
and
occurring as epiphytes on other Algae. G. geminatum (Lyngb.) Ag. is the

largest species (length of valves 90 120 p.; greatest breadth 35 40 p fig. ;

140 A and B) and is principally confined to hilly districts, often forming

thick felt-like masses of a greyish-white colour on dripping rocks.

Genus Rhoicosphenia Grim., 1860. In the valve- view the

frustules are form to those of Gomphonema, being
similar in
fusiform with dissimilar poles in the girdle-view they are cuneate
;

and considerably curved. The valves are dissimilar, the upper

valve only possessing a pseudo-raphe and no nodules, whereas the
lower valve possesses a raphe and central and polar nodules.
R. curvata (Kiitz.) Grun. is common all over Britain, often clothing in
dense masses the thicker branches of species of Cladophora length of valves;

1345 fji.

Family 3. COCCONEMACE.2E.
The family principally characterized by the
is curvature of the
frustules in the plane of the girdle, so that the valve-view always
appears bent or sublimate. The girdle-view is straight and
generally subrectangular. The frustules are symmetrical about a

longitudinal plane and a median transverse plane.

Each valve
possesses a raphe, sometimes straight,
but more often curved, and
situated more or less near the concave margin. There are distinct
There is one large
polar and central nodules except in Epithemia.
in each cell, the median part of which is
parietal chromatophore
disposed within
Two auxospores arise
the concave girdle-face.
without conjugation from two mother-cells, growing side by side
as elongated rejuvenized cells.

There are three British freshwater genera :

A. Valves without transverse ribs.

* Cells not very asymmetrical raphe removed from
;

the edge of the girdle girdle small, without strise

;
Cocconema.
** Cells strongly asymmetrical central nodule very near
;

the girdle on the concave side girdle broad, with

;

longitudinal striations Atnp/tom.

B. Valves with transverse ribs ; raphe often strong and conspicuous.
Epithemia,
 Cocconemacece 299

Genus Cocconema Ehrenb., 1829. [Cy rubella Ag., 1830;

Encyonema Klitz., 1833.] The frustules are asymmetrical, straight,
ormore often sublimate, being strongly
attenuated from the middle towards
the extremities, which are obtuse. In
the valve-view one margin is strongly
convex, whereas the other may be
slightly convex or concave. In the
latter case it is generally tumid in
the middle. The raphe is well-marked
and is nearer the concave or the less
convex side of the valve. The valves
are striated, the striae slightly radiat-

ing from the raphe and consisting in

many species of distinct series of dots.
The girdle- view is straight, often sub-
rectangular, and the actual girdle is

generally very narrow. In some species

the frustules are free, in others they
are stipitate, and in others they are
enclosed in gelatinous tubes, these A

differences in habit having been form-

Fig. 141. A, Cocconema lun-
erly utilized as generic characters. ceolatiun Ehrenb., from Shipley
Glen, W. Yorks. ( x 500). B and
C. lanceolatum Ehrenb. is the largest
ovalis from
C, Amphora Kiitz.,
and one of the most abundant species of Moidart, Inverness (x500).
the genus ;
length 80 150
of valves /j. ;

fig. 141 A. Ehrenb. and C. cymbiforme Ehrenb. are smaller species

C. Cist u la
almost equally abundant. C. Ehrenbergii (Kiitz.) nob. is a large species of
rare occurrence. C. cuspidatum (Kiitz.) nob. is widely distributed and often
abundant on wet rocks in mountainous regions, frequently forming gelatinous
masses of a greyish-brown colour. The frustules of C. prostratum (Berk.)
nob., C. ccespitosum (Kiitz.) nob. and C. gracile (Rabenh.) nob. occur in more
or less linear series enclosed in gelatinous tubes.

Genus Amphora Ehrenb., 1831. The frustules are asym-

metrical, curved or sublimate in the valve-view, with a well-marked
raphe near the concave side. The central nodule is adjacent to
the concave margin and is sometimes widened into a stauros. The
striation of the valves is similar to that of Cocconema. The
girdle-view is elliptical with truncate apices and the girdle is

broad with irregular longitudinal striations. The chromatophore

is similar to that of Cocconema, from which genus Amphora
300 Batillariece

principally differs in the position of the raphe, in the form of the

girdle-view, and in the broad striated girdle.

Most of the species are marine, but A. ovalis Klitz. is a common fresh-
water species with valves 50 70 p in length fig. 141 B and C.
; very A
small variety of it var. pediculus Kiitz. occurs as an epiphyte, generally on
other Diatoms, such as Nitzsckia sigmoidea.

Genus Epithemia Breb., 1838. The frustules are solitaryand

free-floating or often epiphytic on other Alga? and more highly
organised plants. In the
valve-view they are slightly
curved or lunate, with an
inner concave margin and
an outer convex margin.
The valves are
apparently
without a true raphe, which
is replaced by an excentric

pseudo-raphe situated close

to the concave margin. O.
Miiller has shown, however,
that one species possesses a
true raphe, and it is highly
probable that the so-called
'

pseudo-raphe of all the

'

other species is morphologic-

ally a true raphe.

There are
strong transverse ribs in the
valve-view which represent
Fig. 142. A, girdle-view of Epithemia
gibba Kiitz., from Wicken Fen, Cambridge. transverse septa, and the
B, valve-view of E. turgida (Ehrenb.) Kiitz.,
from Keighley, W. Yorks. C, girdle-view valves are also sculptured
of dividing specimen of E. tuniiJa, from with transverse rows of
Lerwick, Shetlands. D, girdle-view of E.
beads or finely punctate
Argus (Ehrenb.) Kiitz., from Wicken Fen,
Cambridge. (All x 450.) stria3. The girdle-view is

subrectangular, lanceolate,
or broadly elliptical with truncate
apices. The transverse septa
also show as costse in the to the convex nature
girdle-view owing
of the valves, and sometimes terminate in small globular
they
expansions at their point of contact with a partial longitudinal
septum, which is situated on each side of the girdle between
it and the valves. There is either one large chromatophore in
 Nitzschiacece 3 1

each cell or two smaller ones. Auxospores are formed in pairs

from two mother-cells after conjugation.
The genus is almost entirely freshwater in habit, E. turgida (Ehrenb.)
150 yx;
Kiitz. being the most abundant British species; length of valves 70
tig. 142 B and C. E. yibba Kiitz. (fig. 142 A) and E. zebra (Ehrenb.) Kiitz.

are also common species in all kinds of localities. E. gibberula Kiitz. var.

producta Grim, and E. Argus (Ehrenb.) Kiitz. var. alpestris (W. Srn.) Eabenh.
often occur in large quantity in mountainous areas.

Sub-order 4. NITZSCHIOIDE^E.
In this sub -order of the Pennatse the frustules are elongated
and asymmetrical, generally with more or less of a sigmoid curva-
ture. Each valve possesses a. keel in the sagittal line, and the
two keels may be diagonally opposite or both displaced to the
same side of the frustule. The edge of the keel is usually furnished
with strong carinal dots. In transverse section the frustules are
rhombic.

Family 1. NITZSCHIACE^.
This the only family of the sub-order. The valve-view is
is

generally straight, with attenuated apices, and a row of carinal

dots, either median or at one edge. The girdle-view is linear or
sigmoid, with truncate apices. The keel is along the sagittal line

of the valve and contains a true raphe. The chromatophores are

variable ;
the contain one diagonal plate-like chro-
cells may
matophore, or two smaller diagonal plates, or a large parietal plate
the median portion of which is opposite one girdle-face.

There are three genera, distinguished as follows :

A. Keel median valves a little convex cells joined

; ;

to form loose bands Bacillaria.

B. Keel displaced to one side valves convex cells free.
; ;

* Keels of two valves Nitzschia.

diagonally opposite
** Keels of two valves
displaced to same side of
frustule Hantzschia.

Genus Bacillaria Gmelin, 1788. The frustules are straight

and united to form plate-like or ribbon-like colonies, which exhibit
a gliding movement of one frustule over another. The girdle-view
with truncate apices and with a row of carinal dots along
is linear,

each lateral margin. The valve-view is narrowly linear, with

attenuated apices and a median row of carinal dots. The valves
302 Batillariece

are slightly convex, transversely striated, and furnished with a

median keel.

B. paradoxa Gniel. occurs in the freshwater dykes and drains of the north-
east arid east of England ; length of valves 60 70 p..

Genus Nitzschia Hassall, 1845; em. Grim., 1880. [Tryblionella

W. Sm., 1853 Grunowia Rabenh., 1864 Nitzsckiella Rabenh.,
; ;

1864.] The frustules are generally curved, rarely straight, usually

free-floating or more rarely forming a thin stratum. Each valve

a keel which is displaced to one side, the two keels being
possesses
displaced to opposite sides of the frustule and therefore diagonally
opposite each other. The valve-view is elongated or elliptic-
lanceolate, with attenuated and capitate or even rostrate apices,
and possesses a row of conspicuous caririal dots at one edge. The
girdle-view is often elongated with

parallel margins
and truncate apices,
but sometimes the sides are inflated
and there is a median constriction.
It is occasionally straight, but more
often sigmoid. The edge of each
keel is furnished with a row of strong
carinal dots, which are sometimes pro-

longed into short ribs, and the valves

are transversely striated.

This large genus of Diatoms has been

divided by Grunow into 22 sections (includ-

ing Bacillaria). There are about 24 British

freshwater species, of which N.palea (Kiitz.)

B W. Sm. (length of valves 2065 p.} and Jf.

sigmoidea (Ehrenb.) W. Sm. (length of valves
up to 480 /x) are the most abundant and at
the same time exhibit the greatest extremes
r

Fig. 143. A and B, two of si/.e. X. linearis (Ag.) W. Sm. and J .

single valves of Nitzm-liin c<ni- communfs Rabenh. are common species, and
xtricta Pritch., from
(Kiitz.)
J T sinuata (W. Sm.) Grun. is general on
Hawksworth, W. Yorks. ( x 500).
.

C and D, N. siumoiiirn (Ehrenb.) dripping rocks. X. aciculm-ix W. Sin. is a

W. Sm. C, short, straight, ab-
; small species with greatly produced ex-
normal form in process of di- it is often exceedingly abundant
tremities;
vision ("irdle-view, x 400) D,
ponds and ditches, and is remarkable
;

in for
transverse section (after Schiitt,
from Pfitzer). the rapidity of its movements.

Genus Hantzschia Grun., 1877. This genus differs principally

from Nitzschia in the position of the keels of the two valves, which
 Surirellacece 303

are displaced to the same side of the frustule. The valve- view is

somewhat curved with rostrate apices, and the girdle-view is

straight and linear-rectangular. The carinal dots are very con-
spicuous.
H. Amphioxys (Ehrenb.) Grun. is the only freshwater species and is a
frequent British Diatom. It often occurs in prodigious quantity on damp
earth length of valves 45 75 /x.
;

Sub-order 5. SURIRELLOIDEJE.
The frustules are generally symmetrical with regard to the

sagittal axis and each valve frequently possesses two rows of wing-
like projections (or alse). There is a median pseudo-raphe without
any trace of nodules, which is sometimes situated on a sagittal
keel. The valves are generally strongly costate.

Family 1. SURIRELLACEJE.
The Diatoms family exhibit considerable variety of
of this
form, although they are almost always symmetrical with regard to
the sagittal axis. In the valve-view they may be elliptical, linear,
ovate, cuneate, or subcircular, with a median pseudo-raphe from
which radiate strong costse (or ribs). In some forms the valves
are winged, each being furnished with wing-like outgrowths which
project outwards at the junction
of the valve and the girdle. The
is sometimes situated on a pronounced sagittal keel.
pseudo-raphe
The valve-faces are occasionally undulate, the undulations showing

plainly on each side of the girdle-view,

and the frustules are
rarelv twisted. There are two large chromatophores in each
cell, one within each valve-face, and sometimes with projecting
internal lobes. In all cases the frustules are solitary and free-
floating. One auxospore is
produced by the conjugation of two
mother-cells.

The family includes the three following genera:

A. Valve-faces undulate r////^//^'''"'"-
B. Valve-faces not undulate.
* Valves
elliptical or keel-shaped, straight,
with strong transverse ril>s; pseudo-raphe
of one valve 'parallel to that of other Surirella.
** Valves circular and
saddle-shaped pseudo- ;

raphe of one valve at right angles to that

( if other- Ca mpylodiscus.
304 Bacillariece

Genus Cymatopleura Turpin, 1827. The frustules are broadly

elliptical or sublinear in the valve-view, often broadly concave at

each side and with subacuminate
apices. The valves are furnished
with a distinct but inconspicuous
pseudo-raphe and with very fine
transverse striations.Along the
margins are short which
costa?
simulate coarse beads, and the
valve-faces are undulate. In the
girdle- view the frustules are linear
with a number of large undula-
tions along the lateral margins.
The genus is
entirely freshwater
in habit.
Fig. 144. A, Cymatopleura ellip-
tica (Breb.) W. Sm., from Comberton, C. Solea (Breb.) W. Sm. is the com-
Cambridge (
x 350). B and C, C.
monest British species the valves are ;

Solea (Breb.) W. Sm. B, girdle-view

elongated and slightly narrowed in the
;

of large specimen from Esholt, W.

Yorks. C, valve-view of small speci-
;
median portion; length 50 130/z;
fig. 144 B and C.
men from Cornwall x 350). (
C. elliptica (Breb.)
W. Sm. is almost equally abundant
and possesses broadly elliptical or elliptic-lanceolate valves; length 80 140 /*;
fig. 144 A.

'

Genus Surirella Turpin, 1827. [Original spelling Suriraya.']

In the valve-view the frustules are elliptical, linear, ovate, or
sometimes twisted. There is a median pseudo-raphe and strong
transverse costa3. The pseudo-raphe of one valve is parallel to
that of the other. The valves possess a sagittal keel and short
wing-like projections (alse) along their margins. The four sets of
ala3 are best seen in a transverse section. The girdle- view is
subrectangular, oblong, or cuneate, and the girdle exhibits irregular
longitudinal striations.
S. biseriata Breb. (length of valves 100 170 p.; fig. 145 A) and robusta .

Ehrenb. (length of valves 160 230 p.) are two of the largest and most
frequent freshwater species. A variety of the latter, S, robusta var. splendida
(Ehrenb.) V. H., is general in the British freshwater plankton (fig. 145 C).
A much smaller species, A', ovalis Brc'b., with numerous varieties, is very
abundant length of valves 16 80 /z. S. spiralis Kiitz. is remarkable for the
;

twisting of the valves round the longitudinal axis.

 Surirellacece 305

Fig. 145. A, Surirella biseriata Breb. from Adel Bog, W. Yorks. (valve view,
,

x 400). B, S. linearis W. Sin., from Mickle Fell, N. Yorks. (girdle view, x 400).
C, S. robiista Elireub. var. splendida (Ebrenb.) V. H., from near Penzance,
Cornwall (valve view, x 400). D and E, Campylodiscus Hibernicus Ehrenb.,
from Baildou, W. Yorks.; D, valve view; E, view showing the saddle-shaped
frustule; x 400.

Genus Campylodiscus Ehrenb., 1841. The valves are ap-

parently irregularly circular, although in reality they are perfectly

circular. The apparent irregularity is due to their curvature, the
frustule Each valve possesses a median
being saddle-shaped.
pseudo-raphe, and the pseudo-raphe of one valve is at right angles
to that of the other. The valves are furnished with costae,

generally short and often beaded. The genus is mostly marine.

C. Echineis Ehrenb. and C. Hibernicus Ehreub. (fig. 145 D and E) are
general but scarce in the freshwaters of the British Islands. Diam. of valves
about 100 LI.

W. A.
Class 6. MYXOPHYCE^] (or Cyanophycese).

THE Myxophyceae (or Blue-green Algse) is the most primitive

class of the Algae, some of the lowest forms having considerable
resemblance to the Bacteria. They are often termed the Cyano-
'

but the earlier name was

'
1

phyceae , Myxophyceae given a' definite

and restricted meaning by Stizenberger, so as to include all those

plants now
recognized as the blue-green Alga?. (Consult Introduc-
tion, page 3.) It also emphasizes the most conspicuous habit of
these plants, namely, the manner in which the great majority of
them exist in gelatinous A few of them are
masses or
strata.

unicellular, some are

colonial, and others filamentous. They are
found everywhere in damp and wet situations, and many of them
are almost entirely subaerial in habit. In moist climates many of
the richest tints of the landscape are due to the presence of Algae
of this class, which occur in every conceivable situation on rocks,
stones, and the trunks of trees, and some regions they give
in
a decided character to the country 2 The filamentous species
.

frequently form compact, felt-like, mucous or leathery patches of

various colours and of considerable extent, occurring on the vertical
faces of rocks which are kept permanently moist or wet. Numerous

Myxophyceas occur in the sea, although they are more abundant in

fresh water. Many of them exist in quantity in both the marine
and freshwater plankton.
Some of the Myxophyceaa have become adapted to a life in hot
water, and they constitute the principal vegetation of hot-springs.
The part played by certain of these Alga? in the formation of rock-
1
There was little necessity for the re-naming of this class by either Babenhorst
(who termed it the Phycochromophycete) or Sachs (who termed it the Cyanophycese).
The class already possessed much the best name that has yet been given it (the
Myxophyceae), and the exactitude of its limitations could not be improved upon
by either of those authors.
The Pedras iici/raa of Angola are due to the prolific growth of ScytoHema
'

uK var. choroi/rajihicuin ride Westcfe G. S. West in Journ. Bot. 1897, p. 303.

;
 307

masses by the extraction of carbonate of lime or silica from the

water of hot-springs is considerable. The deposits formed around
the hot-springs in many parts of the world consist of brightly-
coloured basins or terraces of travertine and sinter. The colours
are very varied, being all shades of yellow, orange-red, pink, blue,
and blue-green, and are due to the presence of brilliantly-coloured
Alga? within the deposit. In the case of the travertine deposits
the deposition of the carbonate of lime is due very largely to the
extraction by the Algae of the carbon dioxide dissolved in the
water.
That Alga? do actually cause the elimination of carbonate of
lime from the water was first shown by Cohn and Weed has
1 2
,

of the Myxophycese in
given a very able account of the assistance
the formation of the travertine and sinter deposits of the Yellow-
stone National Park, U.S.A. He states that from to a -jL of an ^
inch of travertine is formed in three days. He found that the
character and colour of the deposit depended upon the temperature
of the water and the situation of the spring or geyser. The highest
temperature at which filamentous Myxophycese are known to exist
is 85 C.
3
but unicellular Algse have been observed by Brewer in
,

4
California in water at a temperature of 94'5 C.
The cell-wall, which is never absent, is composed partly of
cellulose, partly of pectose compounds, and very largely of chitin.
5
It presents a similarity to the cuticle of higher plants offering ,

considerable resistance to chemical reagents, and it sometimes

contains silica 6 .

Many forms are embedded in a somewhat

of the colonial
extensive mass of mucilage, the external surface of which is
covered with a thin cuticle, and in most of the filamentous forms
there is secreted either a thin mucous sheath or a tough chitinized
sheath, often lamellose in character. This sheath is undoubtedly
secreted by the enclosed cells, and in some genera (e.g. Lyngbya)
all stages are met with
between a thin hyaline investment and a
hard, lamellose, sheathing tube.

1
Cohn in Abhandl. der Schles. Gesellsch. Nat. 1802, p. 35.
Weed, 'Formation of Travertine and Siliceous Sinter by the Vegetation of
2

Hot Springs.' Rep. U.S. Geol. Survey, 1887-8.

3
G. S. West, Some Algas from Hot Springs,' Journ. Bot. July 1902, p. 241.
'

4
W. H. Brewer in Amer. Journ. Science, ser. 2, xli.
5
R. Hegler in Pringsh. Jahrb. fiir wissensch. Bot. 1901, xxxvi.
Hyams & Richards in Technol. Quarterly, Boston, 1902, xv, pp. 308315.
202
308 Myxophycece
The pigment principally phycocyanin, which
of the cells is

partially hides the chlorophyll

and gives many of these plants
their characteristic blue-green colour. In some species, however,
1

pigments of other colours are present, such as carotin, myxophycin

and polycystiri2 In the great majority of the blue-green Algse the
.

is lodged in the cytoplasm in the form of small granules,

pigment
each of which usually contains a mixture of chlorophyll and
phycocyanin. These granules may occupy a somewhat indefinite
extent of the cytoplasm or they may be restricted to certain
regions, and it is a vexed question whether
or not this pigmented

part of the cytoplasm should be regarded as a true chromatophore.

Stockmeyer
3
and Zacharias 4
have each stated that the

pigmented protoplasm cannot be considered as a true chromato-

phore. Massart 5 also considers that although the Coloured layer
functions as a plastid, it cannot be regarded as a true chromato-
the
phore on account of its indefinite limits towards the interior of
cell,and the fact that it may contain both gas and liquid vacuoles.
On the other hand, Deiniga 6 finds in certain blue-green Algse
(such as Aphanizomenon and Nostoc) structures which he regards
as true chromatophores, having the form of a more or less

reticulated or perforated plate in contact with the cell-wall.

Zukal 7 also considers that Tolypotlirix possesses a true chromato-

9
phore, and Fischer and Hegler
8
each regard the pigmented
peripheral layer of protoplasm as a parietal chromatophore.
The
latter carefully described the granular disposition of the pigment
10
and termed the pigmented layer a cyanoplast. Hieronymus
concludes that the pigmented peripheral layer is of the same
nature as the chromatophore of higher plants, but not identical
with it; and Wager 11 also states that its structure recalls the
chromatophore of other organisms. In the family Glaucocystacese
there is a true chromatophore.
I
Chodat in Journ. Bot. de Morot, 189(5, x. Mi/.rophycin is identical with Sorby's
"pink pjiycocyanin."
-
Zopf in Berichte Deutsch. Bot. Ges. 1901, xviii.
Stockmeyer in Berichte Deutsch. Bot. Ges. 1S94, xii.
:1

4
Zacharias in Bot. Zeitung, 1891, xlix ;
in Abhandl. a.d. Geb. Natunv. Ver.
Hamburg, 1900, xvi.
5
Massart in Recueil de 1'Inst. Bot. Univ. de Bruxelles, 1902, v.
"
Deiniga in Bull. Soc. Imp. Nat. Moscou, 1891, no. 2.
7
/ukal in Berichte Deutsch. Bot. Ges. 1892, x.
Fischer, Unters. iiber d. Bau d. Cyanophyceen u. Bakterien, Jena, 1897.
*

y
Hegler in Pringsheim's Jahrb. fur wissensch. Bot. 1901, xxxvi.
10 in Beitriige zur Biol. d. Pflanzen (Colin), 1892, v.
Hieronymus
II
Wager, Eeport Brit. Assoc. 1901 (1902), p. 830.
 Myxophycece 309

The pigment in the cells of the Myxophycese is

unquestionably
variable in its disposition, and although in certain of these plants it

occupies a somewhat indefinite extent of the cytoplasm, in others

itis
undoubtedly collected into definite areas. It must be re-

membered, however, that indefiniteness of the pigmented areas is

not confined to the blue-green Algae alone, but is often observed
in some of the lowly Protococcoidese, and even in such a highly

specialized group as the Desmidiacese.

Kohl has regarded each of these minute granules of pigment
as a chromatophore, but I prefer to adopt Hegler's idea of a

cyanoplast for that part of the protoplasm in which the granules

of pigment are collected. This cyanoplast, which can be regarded
as an archaic type of chromatophore, exhibits much variation in
the degree of its differentiation, and attains in one family the
Glaucocystacea? the highly differentiated condition met with in
the higher groups of Algae. This is one of the reasons which
has induced me to primarily subdivide the Myxophycese into the
Glaucocystidese and the Archiplastidese.
The pigment is generally confined, as mentioned above, to the
more peripheral areas of the protoplasm, the clear central portion
of which often stands out prominently as a rounded mass. This
has received the name of the 'central body' and of late much
attention has been given it, more particularly with the view of

determining whether it should be regarded as a nucleus or not.

'

Most _ of the recent Jnvestigations show that JJiaJj2iiO-tral -body

_

differs considerably in its structure from the cell-nucleus of higher

plants.
Stockmeyer and Zukal have each denied that the 'central body'
has any relation to the nucleus of higher plants, and Marx 1 stated
that he had obtained negative results in his search for a nucleus.
2 ' '

Zacharias states that the term nucleus should not be used for
'

body as nothing is known of the part it plays in the

'
the central

economy of the cell, and Massart affirmed that there was no reason
to consider it as a nucleus as it was sometimes vacuolated and had
no definite outline. Fischer, and also Palla 3 found no chromatin ,

'
'
in the central body of the blue-green Alga? they examined, but
other observers have described the presence of both granules and
1
Marx '
Untersucb. liber d. Zellen d. Oscillarien,' Erlaugen, 1892 (vide Bot.
Centralbl. 1893).
liii,
2
Zacharias in Bot. Zeitung, 1892, 1.
a
Palla in Pringsheim's Jahrb. fur wissensch. Bot. 1893, xxv.
 310 MyxopliycecB
filaments of chromatin. The investigations of Wille 1 Zacharias 2 , ,

and Scott 3clearly show that the cells of the blue-green Algae contain
a body of a nuclear character. Hegler has shown that at least in
some cases, the central body' consists of a faintly stainable ground
'

substance in which is embedded a small quantity of chromatin, but

that it differs from a nucleus in the absencejoLa-jmclearjnembrane
and nucleolus and the still more recent investigations of Wager 4
;

and Kohl 5 have amply confirmed these observations. Dangeard 6

describes a nucleus in Merismopedia, but states there is no trace of
a nucleolus.
Kohl
states that the nucleus of the blue-green Algge not only
from that of higher plants in the absence of a nuclear
differs
membrane and nucleolus, but also in its remarkable form. It
possessesnumerous radiating outgrowths of a pseudopodium-like
which sometimes extend as far as the cell-wall. It may
character,
be that these nuclear outgrowths are in some instances partly
responsible for the indefinite nature of the chromatophore. In
addition to a certain amount of chromatic substance the nucleus
contains a number of granules of albuminous material, which have
been termed '
central granules.' These have not been observed
outside the nucleus.
It would appear from these and other observations, that the
'

body of the Myxophycese differs from a true nucleus in

'
central
certain essential points, such as the absence of the nuclear
membrane and nucleolus, and in its extraordinary form ;
but at
the same time chromatin is
undoubtedly present, and Hegler and
Kohl have each observed a polar separation of the chromatic
/ substance during division, accompanied by the formation of a
chromatic figure. From a consideration of these facts the term
'
'

open nucleus suggested by Hieronymus in contradistinction to

'

the closed nucleus of higher plants, appears to be a very suitable

'

one. Hieronymus was the first to point out the presence of

chromatin and the absence of a nuclear membrane in the so-called
central body,' but he was of the opinion that cell-division was very
'

largely, if not quite, independent

of this structure.

Wille in Berichte Deutscb. Botan. Ges. 1883, p. 243.

1

a
Zacharias, torn. cit. 1885.
Scott in Journ. Linn. Soc. Bot. xxiv, 1887, pp. 188192.
;{

4
Wager in Report Brit. Assoc. 1901 (1902), p. 830.
5
Kohl, Ueber die Organisation und Physiologie der Cyanophy. und die mitot.
'

Teilung ihres Kernes,' Jena (1903).

6
Dangeard in Le Botaniste, 1892, iii.
 MyxophycecB 311

The observations of both Hegler and Kohl seem to show that

(or a substance much resembling it) is the first product
of carbon-assimilation. Chodat has pointed out that mucilage,
soluble starches and cyanophycin may make their appearance in
all Cyanophycin is a reserve albuminous
parts of the cytoplasm.
substance containing both nitrogen and phosphorus, and it occurs
in small granules which swell up rapidly on the addition of hydro-
chloric acid. These granules occur abundantly in the spores of the

Myxophyceas and are used up during their germination.

Minute oil-drops may also occur in the cytoplasm.
1
Etard-and Bouilhac state that Nostoc punctiforme can maintain
itself by a saprophytic existence in absolute darkness, as it has the

power of assimilating organic substances such as glucose.

Certain of the free-floating Myxophycese of the genera
Gloeotrichia, Anabcena, Ccelosphcerium, etc., which sometimes occur

abundantly in the plankton, contain dark red granules scattered

through the cytoplasm. Klebahn and others have asserted that
2

these red granules are gas vacuoles directly concerned with the

floating capacity of the Algas which possess them. This assertion,

however, is by no means proven, Brandt 3
having found them in
species which do not float and there;
is evidence to show that in
some cases they are most probably of an oily nature. Should they
be gas vacuoles, then the cytoplasm of certain
ultimately prove to
of the blue-green Algag contains both gas and liquid (cell-sap)
vacuoles.
In some of the Myxophycese, and possibly in many others, there
is a protoplasmic continuity bi-tween the cells of the filaments.
Wille 4 was the first to point this out in Stigonema compaction var.
brasiliense. Borzi 5 described these protoplasmic connections in
species of Nostoc and Anabo'iia, and Nadson has figured them in
Aphanizornenon and Tolypothrix. It is in certain species of

Stigonema thatprotoplasmic connections are most con-

these

spicuous, these plants presenting a condition precisely analogous

to the protoplasmic continuity of the cells of the Rhodophycese.
In all cases the continuity appears to be effected by a median pore
1
Etard & Bouilhac in Comptes Rendus, cxxvii, 1898, p. 119.
-
Klebahn in Flora, 1895, Ixxx in Bot. Zeitung, 1897, Iv.
;

3
Brandt in Berichte Deutsch. Bot. Ges. 1901, xix.
4
Wille, 'Bidrag til Sydamerik. Algfl.,' Bih. till K. Sv. Vet.-Akad. Handl. 1884.
no. 18. p. 6, t. 1, f. 20.
5
Borzi, 'Le coraun. intracell. della Nostoch.,' Malpighia, i, 1887.
312 Myxophyceoe
through the polar extremities of the cell or through the transverse
cell-walls. This pore is best seen in the young branches of
1
Stigonema ocellatum, and I have previously pointed out that it is
particularly conspicuous if the plants have first been dried and
2
subsequently soaked in water. Fritsch has also described this
protoplasmic continuity between the cells of Anabcena.
In the families Nostocaceae, Scytonemaceas, Stigonemaceas and
Rivulariacese certain special cells, known as heterocysts, occur at
intervals along the filaments. They are sparsely scattered between
the ordinary vegetative cells, and owing to the absence of pig-
mented material they present a very pellucid appearance. Their
walls, which are composed of cellulose, are generally thickened and
of a pale yellow-brown or yellow-green colour. The heterocysts are

frequently larger than the vegetative cells and their walls

commonly possess slight polar thickenings. Each polar thickening
surrounds the apical pore through the cell-wall, this pore being
closed by a minute plate in old heterocysts.

Heterocysts are developed from ordinary vegetative cells,

generally singly from any cell of the filament, or they may be
formed from the two cells contiguous to an existing intercalary
heterocyst. Under natural conditions heterocysts are almost
invariably solitary ingenera except Tolypothrix and Calothrix,
all

but under unfavourable conditions and in cultures they may

become seriate. Both Brandt and Fritsch have described the
occurrence of an intercellular substance excreted during the
formation of heterocysts. This substance, however, can be
frequently observed remote from the heterocysts in Scytonema,
and it is also excreted by the cells of certain species of the Oscilla-

toriacese, a family of blue-green Algae in which heterocysts do not

exist.
The function of heterocysts is not thoroughly understood. They
have been thought to serve as limitations to the length of the
filaments, and they are at times undoubtedly connected with the
breaking of the filaments. In normal plants of the genus Anab<i'n<i
the filaments break readily at all points, and this fracture cannot
therefore be controlled by the heterocysts. Neither does the
structure of a filament of Stigonema support this view, although
in this genus heterocysts limit the hormogones. Hieronymus,
1
West & G. S. West, 'Welw. Afric. Freshw. Alg.,' Journ. Bot. June 1897, p. 242.
2 Fritsch in New Phytologist, iii, April 1904, p. 93.
 Myxophycece 313

Hegler, and later, Fritsch, regard the heterocysts as storehouses

for reserve substances, the latter passing into the heterocyst along
the protoplasmic threads which communicate with the adjoining
cells. It must be remarked, however, that only under very un-
favourable conditions would this function be taken up by the
of a
heterocysts, as under normal circumstances the cell-contents
heterocyst are absolutely homogeneous.
Brandt has observed (in Nostoc commune and .microscopicum)
1
N
the contents of heterocysts set free as gonidia, which have sub-
sequently developed into new filaments. This germination of the
contents of a heterocyst must have taken place under very excep-
tional and abnormal conditions, and it has not been confirmed. It

suggests the possibility that heterocysts are the lingering

and
abortive relics of a type of spore once possessed by certain of the
Myxophyceae, but which long ago ceased to be functional.
The multiplication of the unicellular and colonial blue-green
Alga? is brought about principally by simple cell-fission, division
occurring in every direction of space or in certain directions only.
Definite or indefinite colonies may thus be produced, which dis-
sociate into smaller groups on attaining their maximum size.
Asexual reproduction takes place in a variety of ways. In some
families, such as the Nostocacea?, Stigonemacese and Rivulariaceae,

spherical or cylindrical spores are formed by the rejuvenescence

and
growth in size of certain of the vegetative cells. In the Chama3si-
divides into a number of small
phoniacese the elongated unicell
which are then liberated from the free apex of the cell.
spores
Among the plants of the entire order Hormogoniese reproduction
the formation "^ These are short filaments
occurs by ^J^^moy^ 1

of cells which are set free from the extremities of the vegetative
filaments, and they ultimately develop into new plants. They can
be considered as a primitive type of multicellular gemmae. Repro-
duction may occur in certain species of Nostoc by means of cocci,
2
which are small cells about the same size as the vegetative cells .

They form a scum on the surface of the water, and each one is
capable of producing a colony by simple
cell-division. Reproduction
does not take place in the Myxophycese, although a
by zoogonidia
few motile, blue-green unicells are known to exist.
Sexual reproduction is unknown amongst the blue-green Alga?,
1
Brandt in Berichte Deutsch. Bot. Gesellsch. 1902, xix.
2
Sauvageau in Ann. Sci. Nat. Bot. iii, 1897, p. 367.
314 Myxophycece
but Borzi 1 has observed the formation of some rather remarkable
spores in the genus Anabcena.
A single vegetative cell divides
into two distinct portions which subsequently coalesce, the product

having the characters of a nascent spore.

Many of the Myxophyceas exhibit considerable polymorphism,
passing through a number of diverse states at different periods of
their life-history. Much confusion has arisen with regard to this
polymorphism and many extraordinary statements have in conse-
quence been made concerning the unicellular forms of the blue-

green Alga
1

Itzigsohn,
.
Hansgirg and Wolle have all regarded the
plants of the Chroococcaceae as stages in the development of the
more highly organised blue-green Algae, and the last-mentioned
"
author has gone so far as to state that it is now clearly evident

that all these so-called unicellular plants constitute nothing more

2
or less than conditions in the plant-life of higher forms ." I have

previously pointed out the absurdity of this statement

3
which is ,

erroneously based upon one of the best known facts concerning the
Myxophyceae, namely, the ability of many of the lower forms of
blue-green Algae to live only under the same conditions of environ-
ment as the higher forms with which they are so frequently
intermingled. There is no direct evidence in proof of the generic
or specific identity of many of these forms which live intermingled
in a common gelatinous matrix, and there is rarely much difficulty
in discriininatingbetween the more ^r less unicellular stages of
the higher types and the unicellular or colonial plants of a lower

type.
Some
of the Myxophyceae, principally of the genera Scytonema,

Stiyonema, and Nostoc, are regularly found in symbiotic relation-

ship with Fungi to form the dual organisms known as Lichens.
The Algae which have thus lost their individuality become con-
siderably modified and generally lose almost all traces of their

original specific characters.

A
few blue-green Algae belonging to the Oscillatoriaceae exhibit
spontaneous movements, generally of a slow, oscillating, gliding, or
rotatory character, and concerning which no convincing explana-
tion has yet been offered. It is in the genus Oscillatoria that

1
Borzi in Bull. Soc. Bot. Ital. 1*95, p. 208.
-
Wolle, Freshw. Alg. U.S. p. 330. Plates clxxxiv and cxci in Wolle's text-
book are typical examples of the crude drawings of the Myxophyceas given by that
author.
3
G. S. West in Journ. Bot. Febr. 1899, pp. 52, 53.
 Myxophycece 315

these movements are most conspicuous, and they become more

active under those conditions which cause an increase in the
activity of protoplasm, such as an augmentation of the temperature
It has been stated by Cohn and
l
or of the intensity of the light.

by Correns that the movements only take place when the filaments
2

are in contact with a solid body, and it seems probable that, at all
events in certain species, contact with a solid body or with the
surface film of water is actually necessary for the performance of
these movements. The motion consists of a slow creeping or

gliding of the entire filament which at the same time slowly

rotates around its axis, and this is often accompanied by a slow
oscillation of the extremity of the filament. It must be distinctly
understood, however, that the movements are not of precisely the
same character in all the species of the genus. The filaments of
some species secrete a small quantity of a colourless jelly, and
it was to this secretion of mucilaginous matter that Siebold,

Engelmann, and others attributed the movements. There is,

however, quite as much probability in the explanation put forward
by Hansgirg that the movements are due to osmotic changes.
In Arthrospira the movements consist of a general bending
of the spiral filaments or a slow oscillation of their extremities.
These movements are more vigorous than those of Oscillato ri<i,
spasmodic and jerky, and I have not observed any rotation of the
spiral filaments.
In the genus Spirulina, in which the filament is also twisted
into a close spiral, the movement consists of a rotation around the
axis of the spiral, with a slight propulsion through the water.
In Sp. turfosa Buln., this rotatory motion is relatively rapid.
Many of the Oscillatoriacere emit a very disagreeable odour,
and ponds and ditches which contain quantities of Oscillatoria
often give off bad smells. This is attributed by Jackson and
Ellms 3 to the decay of highly nitrogenous organic matter, in which
partially decomposed sulphur and phosphorus compounds play a
large part.
'

water-bloom and the breaking of the

'

The phenomena of
'

'

meres are due to the sudden and often periodical appearance of

large quantities of a few species of the Myxophyceee, and the

1
Cohn in Archiv fiir mikr. Anat. 11(57, p. 48.
2
Correns in Berichte Deutsch. Bot. Ges. 1896, xiv.
3
Jackson & Ellms in Technul. Quarterly, 1897, x.
316 Myxophycece

Algae concerned in these phenomena are generally species which

normally occur in the plankton of lakes and rivers. The extra-
ordinary rapidity of their increase and the consequent discolour-
ation of the water, together with their equally rapid disappearance,
constitute one of the most remarkable facts in the whole domain
of algological inquiry. Nelson 1 finds that the presence of this
'
'
water-bloom often has a fatal effect on cattle which have been
2
drinking the water .

The Myxophycese, regarded

as a whole, are unquestionably of
a lower type of organisation than any other class of Alga?, and
they must be looked upon as an archaic group which is very little
in advance of the Schizomycetes (or Bacteria).
The group of the Glaucocystidea? is much in advance of the rest
of the blue-green Alga?, the cells possessing a highly specialised

chromatophore and a more highly organised nucleus. This necessi-

tates a primary subdivision of the class into two sub-classes, the

Glaucocystidese and the Archiplastidese.

Sub-class 1. Glaucocystidew. Cells with a distinct and highly
differentiated chromatophore, and with a true
cell-nucleus.

Sub-class 2. Archiplastidece. Cells with a lower type of

chromatophore, often scarcely differentiated, and

with a primitive type of nucleus.

Sub-class 1. GLAUCOCYSTIDE^E.
This sub-class has been instituted in order to include a few
blue-green Alga? which are sharply demarcated from the rest of
the MyxophyceaB by their cytological structure. There is a true
cell-nucleus and also a highly differentiated chromatophore,
both of these characters being a distinct advance on all other
Myxophyceee.

1
N. P. B. Nelson in Minnesota Bot. Studies, 1903, iii, pp. 4750.
-
Other blue-green Algas also appear to be poisonous. Mr Herbert Wright has
forwarded me specimens of Li/nr/bya majuscula Harvey from coral beaches in the
Gulf of Mannar, where it occurs in abundance; and he states that numbers of
horses have frequently been killed by feeding on it.
 Glaucocyxtacece 317

Family 1. GLAUCOCYSTACE^E.
This family includes four genera, only two of which are known
from the British Islands. The plants are unicellular or colonial,
and propagation takes place by the division of the cells in one
direction only, or by division into two, four, or eight cells after a

corresponding division of the nucleus.

Genus Glaucocystis Itzigsohn, 1854. The cells are ellipsoidal,
rarely solitary, but more often occurring in twos, fours, or eights
within the enlarged wall of the mother-cell. In external features
the plants greatly resemble those of the genus Oocystis, but the
chromatophores are central (or axile), and of a bright blue-green
colour. Each chromatophore consists of a central mass with from
10 20 prolongations, which are more or less radiating and con-
siderably curved.
G. Nostochinearum Itzigsh. is widely distributed throughout the British

Islands, but it is somewhat scarce. It is found chiefly among submerged

Sphagnum. Length of cells 13 19 p..

Genus Chroothece Hansgirg, 1884. The cells are ellipsoidal
with a stout enveloping wall, which increases greatly in thickness
at one pole, and is conspicuously lamellose. The chromatophore is
central with radial outgrowths.

Ch. Richterianum Hansg. is known from salt marshes in Bohemia, and a

small form of it (possibly another species) has been observed on wet limestone
rocks in W. Yorkshire. Length of cells (of British form) 20 24 p..

Sub-class 2. ARCHIPLASTIDE^:.
The great majority of the blue-green Algae are included in this
sub-class. The plants are unicellular, colonial, or filamentous, and
in some of the latter (e.g. Stif/onema, Anabcena, etc.), there is a
direct protoplasmic continuity between the cells. The protoplasmic
unit in each cell is of a lower type than in any other of the Algse,
and has been termed by Nadson an 1 '

archiplast.' The nucleus is

of a primitive type, of peculiar form, and without nucleolus 01

nuclear membrane in consequence of the latter character it has
;

been termed by Hieronymus an open nucleus.' The chroma-

'

tophores are likewise of an archaic type, the pigmented protoplasm

being scarcely differentiated and never of any characteristic form.
1
Nadson in Scripta Botanica Horti Univers. Imp. Petropolit. 1895, iv, fasc. 2.
318 MyxophyceoB
There are two orders ofthis sub-class, both of which are

abundantly represented in every part of the world.

Order I.
Hormogonece. Plants filamentous filaments simple
;

or branched, generally
consisting of one or more
rows of cells within a sheath, attached to a sub-
stratum or free-floating.
Order II. Coccogonece. Plants unicellular or colonial com-
monly embedded in a gelatinous matrix, more
rarely
free-floating.

Order I. HORMOGONECE.
This order includes all the filamentous Myxophycese. As a
rule the filaments consist of a simple row of cells, naked or enclosed
within a sheath of varied character. In some of the Oscillatoriacege
and Stigonemacese there are two or more rows of cells contained
within the same sheath. A simple row of cells is known as a
trichome, and the trichome with its enveloping sheath is termed a
filament. In the Scytonemacese, StigonemaceaB, Nostocacea?, and
Rivulariacese heterocysts are more or less
abundantly scattered at
intervals among the vegetative cells, and in some one
genera
extremity of the filament always terminated by a heterocyst.
is

The heterocysts are commonly solitary, but they may occur in

groups, as in some species of Tolypothrios. The filaments are
frequently branched, and there is often a false branch-system due
to the close apposition of the basal extremities of a number of
filaments. New filaments often arise as lateral outgrowths from
the vegetative cells of an older filament. The trichomes of the
newer filaments separate from that of the old filament, but their
sheaths still remain partially fused, and thus a false branch-system
is
produced.
Most of the Alga3 of this order occur attached to a substratum,
and they frequently form dense, felt-like patches or tough gelatinous
masses of various colours. Some of them form more fragile gela-
tinous masses of varied form, either aquatic or on
damp earth, and
others are free-floating.
Asexual reproduction takes place by means of hormogones,
and more rarely by spores.
As a
rule the trichomes are cylindrical with obtuse or narrowed
extremities, but in some of these Algce there is a gradual and
 Stigonemaeece 31 .)
(

conspicuous attenuation of the trichome, either from base to apex

or from the central part towards each extremity. This character
is utilized to subdivide the order.

Sub-order 1. Psilonematece. Trichomes cylindrical, sometimes

narrowed at the extremities.
Sub-order 2. Trichophorece. Trichomes conspicuously attenuated
towards one or both extremities, which are generally piliferous.

Sub-order 1. PSILONEMATE.E.
The majority of the filamentous blue-green Algae belong to
this sub-order. The trichomes are cylindrical or torulose in

character, of a uniform thickness, and with or without a sheath.

The sheath may be hyaline and gelatinous, very thin, or of con-
siderable diameter in many it is tough and lamellose.
;
In some
the sheaths contain no small proportion of cellulose, colouring blue
with chlor-zinc-iodine, but in others they consist entirely of mucus.
The apical cell of the trichome may be obtuse or more or less
conical and attenuated, and occasionally the apical and sub-apical
cells are suddenly
V
attenuated.
The sub-order includes the four following families :

Family 1. Stigonemacecs. Filaments usually stout and branched;

sheaths thick, firm, and often irregular; trichomes with heterocysts,
and frequently consisting of more than one row of cells.
Family 2. Scytonemawte. Filaments with a false branch-system;
sheaths firm and tubular, of more or less equal thickness trichomes ;

consisting of a single row of cells, with heterocysts, but not of uniform

thickness.

Family 3. Nostocac&x. Trichomes commonly tortuose and intricate,

enveloped within a large gelatinous mass, consisting of a single row of

uniform cells (generally torulose), with heterocysts sheaths very deli-
;

cate, mostly confluent.

Family 4. Osi'idiitoriacece. Trichomes consisting of a simple row
of cells, uniform along their entire length except for the apical cells,
which are sometimes attenuated heterocysts absent sheaths variable,
; ;

more or less gelatinous, and sometimes enclosing more than one

trichome.

Family 1. STIGONEMACE.E.
In this family the cells are arranged either in a single series
or in several more or less irregular series within a strong sheath.
This sheath is rarely thin and regular, being more often thick,
320 MyxophyceoB
dark brown in colour, of considerable toughness, and with a very
uneven exterior. The filaments are branched, and the branches
arise by a growth from one of the cells of the main filament. The
filaments increase in length by the division of the cells towards
the apices. The heterocysts are intercalary, being scattered at
intervals between the vegetative cells. When more than one row
of cells exists in a filament the heterocysts are situated in a lateral

position.
of most of the members of this family, with
The robust filaments
their true branches and somewhat irregular outlines, are easily
filaments of the
distinguished from the more uniform and thinner
Scytonemaceae. The presence of more than one row of cells within
the sheath, and the frequent irregular disposition of these cells,
are also distinctive features.
There are only two British genera, Stigonema and Hapalosiphon.
In the former the reproduction normally takes place by the
of the branches,
development of hormogones from the extremities
whereas in the latter the normal method of reproduction is by spores.
Genus Stigonema Ag., 1824. [Sirosiphon Ktitz., 1843.] The
filaments are free-floating or aggregated to form soft, felt-like

.c.j^ioi.

Fig.
3 146. A and B, Stigoncma minutum Hass., from Slieve Donard, Down, Ireland;
A, xlOO; B, x440. C E, St. dcellatum (Dillw.) Thur.,
from Llyn Teyrn,
Snowdon, N. Wales; C, xlOO; D and E, x 440. h, heterocyst.

masses. The cells are always rounded, and are disposed either in
two or more rows, or more rarely in a single The heterocysts
row.
are commonly lateral, or less frequently intercalary. The sheaths
are thick, lamellose, with an uneven exterior, and they are usually
 Stigonemacece 321

of a golden-yellow or brown colour. The branches are generally

short, thick, and irregularly disposed.
or wet rocks, but are
Species of this genus occur principal!}' on damp
sometimes observed free-floating in ponds and lakes. The largest British
species are St. informe Kiitz. and St. mammillosum Ag.,
the filaments of which
reach a thickness of 90 //. Xt. hormoides (Kiitz.) Born. & Flah. is the smallest
known species; thickness of filaments 7 15/t.

Genus Hapalosiphon Nag., 1849. [? Fischerella (Born. &

Flah.) Gomont, 1895.] The filaments are free-floating amongst
other Alga?, or they are rarely
subaerial in moist situations.
The primary filaments are never
very thick, and consist of a single
row (rarely of two rows) of cells
enclosed within a strong sheath
of uniform thickness. The
branches are sometimes of the
same thickness as the primary
filaments, but more often slight-
ly narrower, and they are com-
monly unilateral. They are

mostly long and flexuose, and

are very slightly attenuated.
As a rule they are few and
distant, but occasionally they
arise in unilateral clusters.
The sheaths of the branches
are always thinner than those
Fig. 147. Hapalosiphon Hih,>rni<-us
of the primary filaments, and West &G. S. West, from Glen Caragh,
Kerry, Ireland x 4-10). E, portion of a
generally quite colourless. The
(

row of spores within the sheath.

cell of the primary filament

opposite the base of a branch usually projects into it, and the
cells ofthe branches are proportionately much longer than those
of the primary filament. The spores are formed from the ordinary
vegetative cells, and possess thick yellowish-brown cell-walls.
Often the majority (or even all) the cells of both primary filaments
and branches of some portions of the plant become converted into
1
spores .

1
West& G. S. West in Journ. Bot. June 1897, p. 241.

w. A. 21
322 MyxophycecB
The plants of this genus are most frequently met with in the British
Islands amongst Utricularia minor and submerged Sphagnum. The two
most widely distributed species are H. intricatus West (diatn. of filaments
4 1 p.) and H. HibernicusWest & G. S. West (diam. of filaments 6 10 p;
fig- 147).

Family 2. SCYTONEMACE^.
The plants of this family are at once distinguished from all
others of the Hormogoneae' by their type of branching. Except
in the three rare genera, Desmonema, Hydrocoryne and Diplocolon,
the cells are always disposed in a single series within a strong
tubular sheath of regular thickness. The trichomes of the
primary filament perforate the sheath at intervals- and issue as
long flexuose branches, which develop a sheath of their own.
These false branches arise either singly or in pairs. The trichomes
are cylindrical, but towards the growing end of the filament they
increase in diameter, the cells becoming much shorter and more
rounded. The sheath may be homogeneous and colourless, or
lamellose and of a yellow or brown colour.
The normal reproduction is by hormogones, but in most of the
genera ellipsoidal or globular spores are sometimes produced.
The five British genera can be distinguished as follows :

A. With one trichome in each sheath.

* Filaments
simple, unbranched Microchcete.
* Filaments with false branches
(" pseudo-rami ").
t Branches geminate, arising between the
heterocysts Scytonema.
tt Branches arising singly in the region of
the heterocysts Tolypothrix.
B. With 2 6 trichomes within each sheath.
* Filaments Desmonema.
straight, with basal heterocysts ...

** Filaments contorted within a common sheath

Diplocolon.

Genus Microchsete Thuret, 1875. [Coleospertnum Kirchner,

1878.] The filaments are fixed at the base, erect, or tortuose-
rlexuose, and without branches. The plants greatly resemble
unbranched species of Sci/taiu'ina, and they o'ccur in radiating
tufts <>r soft felt-like masses. Kirchner 1
has placed this genus
in the Nostocaceae, but the presence of a prominent tough sheath
round each individual trichome, and the thick growing apices with

1
Kircbner in Engler and Prantl's Natiirl. Ptianzeiifam. i, 1 a, p. 76.
 Scytonemacece 323

shorter cells, are characters which indicate a clear affinity with

the Scytonemacea3.

J/. diploaiphon Gorn. var. Cumbn'ca'West is known from the English Lake
District ;
thickness of internal sheath 6'5 9 /z. Species of this genus are
very rarely met with.

Genus Scytonema Ag., 1824. [Petalonema Berkeley, 1833 ;

Schizosiphoii Kiitz., 1843 (in part); Symphyosiphon Kiitz., 1843 (in

part) Arthrosiphon Kiitz., 1845.] The filaments generally occur in
;

a dense intricate mass, and are at once characterized by their false

branches. These pseudo-branches arise in pairs between the
heterocysts, each pseudo-branch being a continuation of the
original trichome which has perforated its old sheath. The sheath
is
tough, lamellose, and of a golden-yellow or brown colour it is ;

generally of even thickness, but in a few species (e.g. S. alatuni) in

which it is of great width, the margins are irregular.

Fig. 148. A D, Scytonema mirnliile (Dillw.) Thur., from Cronkley Fell, N. Yorks.
( x 440) C, apex of a ''branch'; D, organ of attachment at base of filament.
;

E, Tolypothrix l/ui/ita (Desv.) Wartm., from Riccall Common, E Yorks

(x440).

M>/ochr<yiix Ag. and >$'. mirabile (Dillw.) Thur. [ = s fi<jur<t.tuin Ag.] are
.S'. > '.

the two most abundant British species, the former often occurring on wet
rocks in large felt-like masses an inch in thickness thickness of fil. 18 36 p.
; ;

thickness of trichomes 6 12/*. S. alatuin (Berk.) Borzi is frequent on wet

rocks of carboniferous limestone, occurring as a brownish-green stratum
which is thickly encrusted with lime ;
thickness of fil. 24 66 // ; thickness of
trichomes 9 15 p..

212
324 Myxdphycece
Genus Tolypothrix Kiitz., 1843. [Indus. Hassallia Berkeley,
1845.] The filaments of this genus are very like those of Scytonema,
and the plants are primarily distinguished by the single pseudo-
branches. The latter are never geminate as in Scytonema and
they always issue in the region of the heterocysts. Both terrestrial
and aquatic species occur, and the sheaths, which are usually
thinner than those of Scytonema, may be either flexible or more or
less fragile. Hassallia could only be separated from Tobjpothrix
by the fragility of its sheaths, and that character is totally in-
adequate as a generic distinction The heterocysts are sometimes
1
.

3-, 4-, or 5-seriate.

T. lanata (Desv.) Wartm. (thickness of fil. 9 12'5 /z ; thickness of

trichomes 7 "5 10 fj.; fig. 148 E) and T. tennis Kiitz. (thickness of fil. 8 10 /A ;

thickness of trichomes 6 8 /*) are the most frequent British species, occurring
amongst various aquatic plants in ponds and lakes.

Genus Desmonema Berkeley & Thwaites, 1849. The fila-

ments of this genus exhibit a slight dichotomous branching,
and two or more trichomes are included in one sheath. The
heterocysts are only found at the base of the filaments.
D. Wrangelii (Ag.) Born. & Flah. is a very rare British Alga, occurring in
penicillate tufts 5 6 mm. high thickness of trichomes 9 10 /*.
;

Genus Diplocolon Nag., 18-57. The filaments possess single

or geminate pseudo-branches of a similar nature to those of
Scytonema but much shorter and greatly contorted. They consist
of one trichome within a sheath, and a number of filaments and

pseudo-branches are included within a larger common sheath.

D. Heppii Nag. is known from Yorkshire. It forms a brownish-green

gelatinous stratum on damp calcareous rocks; thickness of fil. 20 28 p.;
thickness of trichomes 6 10 j..

Family 3. NOSTOCACE^.
The plants
of the Nostocaceae are of a simpler type than

any others of the Psilonemateae which possess heterocysts. The

1laments are never fixed by one extremity and they are never
1

branched. The trichomes consist of a single series of cells of

uniform character. The cells are frequently torulose and the

1
West & (T. S. West in Journ. Bot. July 1897, p. 266.
 NostocacecB 325

heterocysts may be terminal or intercalary. The sheaths are very

delicate and invariably gelatinous in character. In some they are
more or less distinct, of variable diameter and closely adhering to
the trichomes; but in others they become confluent, forming a
watery mass of jelly which encloses a large number of trichomes.
It is in the genus Nostoc that this gelatinous mass becomes most

conspicuous. Whenever there is a fracture of a filament the

trichome and the sheath both break at the same point, and empty
sheaths, such as are commonly found in the Stigonemacea? and
Scytonemacese, are never met with. The trichomes are sometimes

straight and rigid, but they may be contorted to form densely

interwoven masses.
Reproduction occurs by the formation of hormogones and
spores. The former are produced by the fragmentation of fully
grown trichomes, and in some genera they are set free by the
dissolution of the enveloping mucilage. Spores of a spherical,
ellipsoidal, or cylindrical form are developed from the vegetative
cells, and invariably arise in relation to the position of the hetero-

cysts. They may be developed singly or in series, and by reason

of their definite form and position they furnish a character of

primary importance in the discrimination of genera and species.

About half the species of Nostoc and Cylindrospermum are
terrestrial in habit, but all the other members of the family are

aquatic. species of Nostoc, and also of Andbcena, are

Some
endophytic in various parts of such plants as Azolla, Cycas, Lemna,
Sphagnum, Anthoceros, etc., and other species of Nostoc are con-
stituents of the thallus of Lichens, especially those of the
many
CollemaceaB. Many species of Anabcena are a marked feature of
the freshwater plankton, and certain species of Nostoc are used as
food by the inhabitants of divers parts of the world.
A. Trichomes flexuose and contorted within a defi-

nite gelatinous investment Nostoc.

B. Trichomes more or less straight, free, or forming

a thinmucous stratum.
*
Heterocysts and spores intercalary.
t Trichomes nude, or with a thin mucous
sheath, free, or aggregated without
order to form a flocculent mass cells ;

equal or longer than their diameter;

spores solitary, geminate, or in short
series -1 ndbaena.
326 Myxophycece
ft Trichomes short, aggregated in parallel
bundles to form thin, feathery, plate-
likemasses Aphanizomenon.
itt Trichomes free; spores seriate, remote
from the heterocysts cells shorter ;

than their diameter Nodularin.

**
Heterocysts terminal and the spores always
contiguous to them Cylindrospermmn.

Genus Nostoc Vaucher, 1803. The plants of this genus are

found as gelatinous masses of a rounded expanded character, or
sometimes attached to a
substratum, at other times
floating freely in the water.
When young the plants
are almost always globose
or ellipsoid, but with ad-

vancing age many species

become exfoliated to form
flattened expansions, often
with lacerated margins.
The trichomes are much
contorted, forming an in-
tricate mass within the

gelatinous envelope, and

they are generally much
denser towards the ex-
Fig. 141). A and B, Nostoc Liiickia Bornet,
from Ben Lawers, Perthshire; A, nat. size; terior. The sheaths are
B, small portion of thallus, x 340. C, N.
never very distinct, and
cieruleinn Lyngbye, from B. Wharf e at Ilkley,
W. Yorks., nat. size. more often they have
become completely merged
with the enveloping jelly. The cells may be spherical, barrel-
shaped, or cylindrical, and the heterocysts are either intercalary,
or in the young forms, terminal. The spores are spherical or
oblong, and are developed centrifugally in series between the
1
heterocysts .

Young Nostocs of different species greatly resemble each other.

They consist of a variously contorted trichome within a relatively
small, elongated, mucous investment, terminated at each end by a
heterocyst.

1
Borzi in Gioruale Botan. Ital. 1878, x, p. 241.
 Nostocacece 327

There are about 15 British species of jVostoc, some of which are much
more abundant than others. N. piscinale Kiitz., N. coeruleum Lyngbye
(fig. 149 C) and N. pruniforme Ag.
are widely distributed in ponds and
ditches, generally occurring as free-floating masses. N. commune Vauch.
prefers damp ground which is frequently inundated. N. muscorum Ag.,
N. humifusum Garni, and N. macrosporum Menegh. are found principally
among mosses on wet rocks. N. sphcericum Vauch. and N. verrucosum Vauch.
commonly occur attached to the rocks and stones in the beds of rapid streams
and rivers. The cells of N. humifusum are only 2 2'5 p. in diameter, whereas
those of JV. macrosporum reach a diameter of 8 9 p.

Genus Anabaena Bory, 1822. [Sphwrozyga Ag., 1827 ;

Tri-
ch or mus Allman, 1842; Dolichospermum Thwaites, 1850.] The
trichomes are straight or circinate, often destitute of all traces of
a sheath, ami are either free-floating or aggregated to form a thin
mucous stratum. The cells are generally of equal size, but some-
times there is a slight attenuation of the trichome, and the apical
cell may be conical. The cell-contents may be either homogeneous
or granular. The heterocysts are numerous and intercalary. The
spores are variously disposed in ; some they are solitary, in others
there are two spores one on each side of a heterocyst, and rarely
they are in short concatenate series. In most species they
arise

centrifugally with regard to the heterocysts, but in some they are

developed centripetally. The latter were formerly placed in another
'genus Sphcerozyga.
As a Anabcenas cannot be kept alive very long after
rule,
collection, unless placed in a large volume
of water. One or two
in a small vessel is usually sufficient to cause a
days glass
disarticulation of the filaments of most of the species, and under
these conditions they often lose their specific characters, develop-
ing forms which are quite unknown in their
natural state.
this genus with
Hansgirg at one time suggested the union of
the genus Nostoc, but there are many good reasons for keeping the
Species of Anabcena
even when found in a
genera separate.
mucous stratum, which is but rarely, never approach in consistency
the tough thallus of a Nostoc with one or two exceptions the
;

trichomes are and

rigid
and they are never at any time so
fragile,
contorted as those of a Nostoc; the spores are generally much more
in the latter genus, and the habits
elongate than those produced
of the two genera are mostly quite different. I doubt if the merest
novice at the study of freshwater Alga? would confound an Anabu'tm
with a Nostoc. He certainly would not had he collected them
328 Myxophycece
himself. Why, under these circumstances, should the genus Nostoc
be made more confusing than it is already ?
There are about 10 British species of the genus, some of which are
relatively abundant in the waters of ponds and lakes. .1. Flos-aqiux Breb.
and A. drcinalis Rabenb. are constituents of the freshwater plankton. A.
oscillarioides Bory and A. incequalis (Kiitz.) Born. & Flah. (fig. 150
A D) are
found among other Algae in still water. Several species occur among Sphag-
num in bogs, but spores seem to be rarely developed in these situations. The
trichomes varyin thickness in the different species from 4 to 10 /*, and the

cylindrical type of spore may reach a length of 60 p..

Genus Aphanizomenon Morren, 1838. The trichomes are

straight and without

a defi-

nite sheath, being aggluti-

nated to form spindle-shaped
or small plate-like bundles,
which float freely in the water.
Each trichome is slightly at-
tenuated towards the extremi-
ties. The spores are solitary,
cylindrical,
much elongated,
and only developed sparingly
between the heterocysts.
A. Flos-aquce (L.) Ralfs is

found floating in the still waters

of ponds and lakes, sometimes in
abundance. The trichomes are
rigid and 5 6 p.
in diameter.
The cells are subquadrate and
the heterocysts are cylindrical.
The spores are /*
6080
in length
Fig. 150. A D, Anab<e,na incequalis and only 7 8 /x in breadth.
(Kiitz.)Born. & Flah., from Epping Forest,
Essex. E G, Cylindrospermum stannalc
Genus Nodularia Mer-
(Kiitz.) Born. & Flah.,
from Eibblehead,
W. Yorks. H, Nodular ia spluerocarpa Born. tens, 1822. The filaments
& Flah., from near Ely, Cambridgeshire.
are free-floating and are
(All x480.)

generally furnished with a

distinct sheath. The hyaline and mucous, closely en-
latter is

veloping the trichome, and sometimes becomes diffluent. The

trichomes are more or less straight, with short, depressed, often
discoidal cells. The heterocysts are also depressed. The spores
are spherical or discoidal, and are developed in series- between the
heterocysts.
 OscUlatoriacece 320

There are three British species of the genus occurring in fresh water (or
sometimes in slightly brackish water). None of them are abundant, although
JV. spumigena Mertens is not uncommon (diameter of trichomes 10 15 p.}.
N. sphcerocarpa Born. & Flah. is known from Cambridgeshire (diameter of
trichomes 67 /x ; fig. 150 H).

Genus Cylindrospermum Kiitz., 1843. The trichomes in this

genus are relatively short, destitute of a sheath, and are mostly

aggregated to form an expanded mucous stratum. The cells are
cylindrical, longer than their diameter, and one end of the trichome
is terminated by a solitary heterocyst. The spores are always
developed from the cell (or cells) next the heterocyst they are ;

generally solitary, but in one species they are seriate. The terminal
position of the heterocysts and spores at once distinguishes this
genus from all the others of the Nostocacese.
There are three British species. C. majus Kiitz. ( = C. macrospermum
Rabenh.) is common on damp stones and earth, often forming a thin stratum
on garden paths or at the edge of a pond thickness of trichomes 5 6 p..
;

C. stagnale (Kiitz.) Born. & Flah. is not uncommon in boggy ditches, particu-

larly in peaty ditches; thickness of trichomes 3'8

4-5 fig. 150 E G.
/j. ;

C. catena-turn Ralfs is a much rarer species occurring on the damp mud of

stagnant rivers and ditches.

Family 4. OSCILLATORIACE^I.
This is largest family of the Psilonematea^
the and is
at once from all the others by the entire absence
distinguished
of heterocysts. The trichomes consist of a simple and uniform
row of cells, which sometimes exhibits a slight attenuation at the
apex. The apical cell may be rounded, conical, or subcapitate,
and
itoccasionally carries a slightly thickened hood or calyptra.
The
trichomes are straight or flexuose, occurring either free-floating or
forming a thin mucous stratum. The slimy stratum formed by
filaments of this family may occur on damp surfaces, on submerged
stones, or on the muddy bottom of ponds and ditches. Some
species of Oscillatoria rise to the surfaces of ponds as brown or

brownish-green flocculent masses, which are often very conspicuous.

This generally occurs in bright sunshine, and the mass floats owing
to the retention of numerous bubbles of oxygen amongst the
intricate network of trichomes. The cells of some species of this
family are joined so closely that the trichome appears perfectly
cylindrical with straight margins.
In other species there is a faint
330 MyxophycecB
between the
constriction cells, or the latter may even be separated
by a mucous layer. In rare instances the trichomes are spirally
twisted.
Sheaths are not always present in this family, and when
present, they are of very varied character. In some genera, such
as Microcoleus and Phormidium, the sheath is always colourless,

transparent, and more or less evanescent, whereas in others, such

as Schizothrixand Plectonema, it is of greater firmness, generally
branched, and often lamellose.
Slow, gliding or rotatory movements are exhibited by some
species of Oscillatoria, Phormidium, Arthrospira and Spindina.
The family issubdivided into two groups :

Sub-family I. Vaginariece. Several trichomes included within a

single sheath, which is frequently branched.

Sub-family II. Lyngtyecx. Never more than one trichome within

a sheath.

Sub-family I. VAGINARIE.E.
This sub-family contains the most highly organised of those
filamentous blue-green Algse which do not possess heterocysts.
The most important feature of the group is the inclusion of two
or many trichomes within the same sheath. The sheaths often
exhibit more
or less branching, and are of two distinct types. In
one type they are firm and lamellose, often brown, brownish-red,
or even blue in colour in the other type they are mucous and
;

hyaline. Those of the

first type
always become of a blue colour
on the addition of chlor-zinc-iodine, whereas those of the second
type very rarely become coloured by this treatment.
A. Sheaths generally coloured ; trichomes loosely aggregated within the
sheath, not very numerous.
* Sheaths
firm, lamellose, hyaline or coloured; con-
taining few or many trichomes Xchizothrix.
** Sheaths
wide, hyaline or yellow-brown, diffluent ;

containing very few trichomes Dasygloea.

B. Sheaths always hyaline, not lamellose, containing a
large number of trichomes Microcoleus.

Genus Schizothrix Kiitz., 1843. [Indus. Inactis Klttz., 1843;

Hyplueothrix Kiitz., 1843 (in part).] The filaments of this genus are
variously agglomerated to form small cushion-like masses, erect
tufts, or a flat stratum, and occasionally they occur floating freely in
 Oscillator iacew 331

water. In some instances there

is a deposition of lime in the

stratum. The filaments

are either simple or variously branched,
and are sometimes of great length. The sheaths are firm and
lamellose, generally attenuated towards the apex, and rarely
colourless. The inner layers of the sheath may be any shade
of yellow, yellow-brown, purple, or blue, and they are usually of
a deeper colour than the outer layers. There are relatively few
trichomes (sometimes not more than two) within each sheath, laxly
disposed, and sometimes spirally twisted.

Fig. 151. A, Schizothrix Miillerii Nag., from Penyghent, W. Yorks. B, ,S'. lardacea
(Ces.)Gom., from Arncliffe, W. Yorks. C, Daxi/fihea nmorpha Berk, (a small
form) from Thursley Common, Surrey. (All x 460.)

The Miillerii Xiig. (thickness of trichomes

largest British species is
*$'.

13 151 and the smallest is S'. d<'Ucut<xint W. & G. S. West

7 fj.; fig. A)
(thickness of trichonies 0'6 O'8/i). S. calcicola (Ag.) Gom. and >$'. lardacea
(Ces.) Gom. (fig. 151 B) are not infrequent on
wet rocks. In S'. funalis W. &
332 Myxophycece
G. West, which is another very small species (thickness of trichomes
S.

7 p.), the branches of the filaments are twisted round each other like
-

0'5
the strands of a rope.

Genus Dasygloea Thwaites, 1848. This genus differs principally

from Schizothriac in the of relatively wider sheath *, which
1

possession
enclose fewer and more remote trichomes.

D. (tmorpha Berk, occurs in permanent bogs thickness of trichomes

;

4 6 p..
A figure is given of a small form of the species (fig. 151 C).

Genus Microcoleus Desmazieres, 1823. [Cthonoblastus Ktitz.,

1843.] The filaments are simple, terrestrial or aquatic in habit,
and are furnished with a conspicuous hyaline sheath. This sheath
is more or less cylindrical,way lamellose, and its apex
not in any
is
generally diffluent. The trichomes
are numerous, closely aggre-

gated within the central part of the sheath, and often spirally
interwoven. The apical cells are acute or acutely conical.
Plants, of this genus are very rare in Britain. M. subtondosus (Breb.)
Gom. (thickness of trichomes 4 5 p.) and M. delicatulus W. & G. S. West
(thickness of trichomes 1-5 2 p. ; fig. 152 A) are known from Scotland.

Sub-family II. LYNGBYE^.

In this sub-family of the Oscillatoriacese there is never more
than one trichome within a sheath, and the latter, although firm
and lamellose in some genera, is indefinite or wanting in others.
The trichomes vary much in thickness and in the relative length
of their cells. As a rule, the thicker the trichome, the shorter the
cells. In Artkrospira, Spirulina, and some species of Lyngbya,
the trichomes are regularly and spirally twisted. In Spirulina
the trichome is unsegmented, thus consisting of a single cell This
genus, therefore, approaches certain genera of Bacteria, such as
Spirillum and Spirocluvte. The extremities of the trichomes may
be obtuse, capitate, acuminate, or even greatly attenuated. Move-
ments of a gliding or rotatory nature are exhibited in the genera
Oscillatoria, Phormidium, Arthrospira and Spirulina.
A. Trichomes consisting of many cells.
* Filaments
simple, or falsely branched ;
sheaths firm ; apices of
trichomes straight,
t Filaments free or forming felt-like masses,
branched pseudo-branches often geminate
;
Plectonema.
ft Filaments forming erect tufts, often branched ;

pseudo-branches solitary Symploca.

 Oscillatoriacece 333

ttt Filaments free and simple, free-floating or

forming a matted stratum Lyngbya.
** Filaments
simple; sheaths thin, always hyaline, mucous, and
more or less diffluent, in some species absent apices of ;

trichomes commonly curved,

t Trichomes more or less agglutinated by their
mucous sheaths ;
cells of trichome often

slightly separated by a thin mucous layer... Phormidium.

tt Trichomes destitute of sheaths, free, straight,
or with curved extremities Oscillatoria.
ttt Trichomes destitute of sheaths, twisted into
a regular spiral Arthrospira.
B. Trichomes consisting of one cell, twisted into a regular
^ nl inn.
spiral fiir

Genus Plectonema Thuret,

1875. The filaments are free-
forming csespitose masses,
floating,
or they are intricately matted
amongst damp mosses and on wet
rocks, forming felt-like or mat-
like expansions. The sheaths are
firm, hyaline or lamellose, and
rarely of a yellow-brown colour.
The trichomes fit closely within
their sheaths, 'and there is often
a slight constriction between the
cells. The filaments give origin
to false branches exactly similar
to those of Scytonema in fact, ;

Plectonema only differs from Scy-

tonema in the absence of hetero-
cysts and in the more irregular
manner of its false branching.

P. Tomasinianum (Kiitz.) Born, is

the largest and most frequent British
species, occurring as greenish-brown
felt-like masses on wet rocks. Thick-
ness of filaments 16 27 n; thickness
of trichomes 12 -5 22 /x length of
;
Fig. 152.A, Microcoleus delicatulus
Thick-walled resting
W. & G. S. West, from Glen Tummel,
cells 3 9 fj..
Perthshire x 350). B ami C, Si/niplt>cii
(

cells, probably of the nature of spores, iniirit/i* from Frizinshall,

Kiitz., W.
have been observed in this species. Yorks. (B, nat. size; C, x 350).
334 MyxophycecB
Genus Symploca Klitz., 1843. In this genus the filaments
are densely interwoven to form a creeping stratum from which
arise numerous erect tufts of variable height. false branching A
is exhibited by the filaments of some species, the branches arising
singly. The sheaths are thin, hyaline, and generally firm; they
partially coalesce in the erect tufts. Only one trichome is present
within a sheath and its apex is straight.
S. muralis Kiitz. is known from W. Yorkshire. It occurs on damp earth,
walls, or trunks of trees; thickness of trichomes 3'4 152 B and C.
4/j,; fig.

Other species occur among mosses and dead leaves, and others in hot springs,
the genus having a world-wide distribution.

Genus Lyngbya C.Ag., 1824. [Leibleinia Endlicher, 1836 ;

Leptothrix Klitz., 1843

(in part) ; Spirocoleus
Mobius, 1889 (in part).]
The filaments are un-
branched, free-floating or
densely intricate, forming
a floccose mass or an
expanded stratum. The
sheaths are firm, of vari-
able thickness, and some-
times lamellose. They
are generally hyaline, but
in a few species they
Fig. 153. A, Lyngbya major Menegh., from
Wimpole Park, Cambridge. B and C, L. ceru- become of a yellow-brown
Gom., from Bradford, W.
gineo-ccerulea (Kiitz.) colour. The trichomes
Yorks. D, Pkormidium iiiolle (Kiitz.) Gom. (a
narrow form), from Bradford, W. Yorks. E and are either obtuse or
F, Ph. tenue (Menegh.) Gom., from Wicken Fen, attenuated at
slightly
Cambridge. (All x 460.)
the apices, and sometimes
there are evident constrictions between the cells.

Species of this genus are widely distributed and occur in very varied
habitats. Those belonging to the subgenus Leibleinia are entirely marine
and live as epiphytes on other larger Algse. There are about 10 British
freshwater species, of which L. Martensiana Menegh. (thickness of trichomes
6 10 fi) and L. cerugineo-coerulea (Kiitz.) Gom. (thickness of trichomes
46 /u fig. lf)3 B and C) are the most
;
abundant. L. ochracca (Kiitz.) Thur.
occurs in water containing much iron, the oxide of iron ultimately forming a
thick deposit round the sheaths of the filaments. L. cestuarii (Mert.) Liebman
is the largest British species which occurs in fresh water (thickness of tri-

chomes up to 24 p.), although it is principally brackish or marine in habit.

 OscillatoriacecB 335

Genus Phormidium Kittz., 1843. [Hyphceothrix Kutz., 1843

(in part); Leptothrix Klitz., 1843 (in part).] This genus is inter-
mediate in character between Lyngbya and Oscillatoria, and
undoubtedly serves a useful purpose for the reception of species
which cannot be strictly referred to either of those genera. The
filaments are simple and invested by delicate hyaline sheaths,
which frequently become confluent or altogether diffluent. The
filaments are often agglutinated to form an expanded stratum,
either on damp earth, wet rocks, or entirely submerged. Some-
times this stratum becomes hard and leathery, but it is more
often soft and slimy. The trichomes are cylindrical, and there is
frequently a constriction between the cells, a feature which is so
prominent in some species as to give the trichome a moniliform or
torulose appearance. The apices of the trichomes may be straight
or curved, and the apical cells are attenuated, capitate, or even
thickened at the extremity.
Plants of this genus are amongst the commonest of the blue-green Algae,
occurring in all kinds of damp and wet situations, and often giving a decided
tint to large areas of damp ground, vertical limestone rocks, or those rocks
and stones subject to the spray of waterfalls. There are some 13 British
species occurring in damp Ph. autumnale (Ag.)
situations or in fresh water.
Gom. (diam. trich. 4 abundant on damp earth, and Ph. tenue
7 /*)
is

(Menegh.) Gom. (diam. trich. 1 2/^i; fig. 153 E and F) is frequent among
other Algee in ponds, ditches, and rivers. Ph. purpurascens (Kutz.) Gom.
forms reddish-purple patches on the vertical faces of wet limestone rocks
(diam. trich. 1'5 2 p.).

[Note: Clonothrix gracillima W. & G. S. West is one of the Schizotny-

cetes of the geuus Cladothrix.~\

Genus Oscillatoria Vaucher, 1803. [Oscillaria Bosc, 1800 '.]

The trichomes are free and cylindrical, without a sheath, or with
one so thin as to be almost imperceptible. Faint constrictions are
sometimes evident between the cells, but more often the edges of
the filaments present a continuous and unbroken line. The cells
vary much in relative length, but in the larger species they are
always much shorter than their diameter. The extremity of the
trichome may be straight or curved, and it is often attenuated.
The apical cell is sometimes much attenuated, and may be capitate,
being frequently furnished with a terminal thickening of the cell-
wall known as a calyptra. The genus is found in great profusion
For reasons for adopting the generic name " Oscillutoria "
1
consult Gomont in
Journ. Botan. Morot, v, 1891, p. 273.
336 Myxophycece
in all kinds of wet situations, and sometimes on damp ground, or
in hot springs. Flocculent masses
of Oscillatoria are often found float-

ing on the surfaces of ponds and

ditches after sunshine, and they

commonly give off a more or less

m,,
r.--.
distinctive odour. The movements
exhibited by the trichomes of this
L
genus are discussed on page 315.
There are about 20 British species of
the genus, one of which (0. decolorata
G. S. West) is destitute of pigment and
lives a saprophytic existence in stagnant
ditches. 0. princeps Vauch. is the largest
i 50
species (thickness of trichomes 25 p.)

and 0. angustissima W. & G. S. West is

the narrowest (thickness of trichomes

Fig. 154. A, Oscillatoria limosa
Ag., from Wimbledon Common, 0'6 /*).
0. limosa Vauch. (thickness of
Surrey. B, 0. irrigiia Kiitz., from trichomes 12 17 154 A) and 0.
p. ; fig.
Shipley Glen, W. Yorks. C, 0.
tenuis Ag. (thickness of trichomes 4
teniiix from Sheep's Green,
Ag.,
Cambridge. D, O. splfndida Grev. 10 p. fig. 154 C) are the most abundant
;

W. & G. S. West,
var. attenitata
species. 0. irrigua Kiitz. (fig. 154 B) is
from Baildon Moor, W. Yorks. common
from
in quickly running water.
E, 0. acuminata Gom.,
Sheep's Green, Cambridge. (All Genus Arthrospira Stizenberger,
x 460.)
1852. The trichomes are cylindrical,
commonly devoid of a sheath, and twisted into a regular spiral.
The latter character is the only distinction from Oscillatoria.
The cells are generally shorter than their diameter.
Jenneri (Hass.) Stizenb. is a rare Alga, which occurs in stagnant
A.
water, or forms a dark-green mucous stratum in trickling water diain. of ;

trichomes 5 8 p. diam. of spiral whorls 9 15 p..

;

Genus Spirulina Turpin, 1827. The trichomes are very narrow

and are twisted into a regular spiral.
They consist of a single elongated cell,
sometimes of great length, an4 without
B
transverse septa. This feature at once
Fig. 155. A. pirulina
hirl'ond Cram., from Lough distinguishes the genus from Arthrospira.
Nciigh, Ireland. B, .S'/*.
$. major Kiitz. is frequent in stagnant water
major Kiitz., from Wicken
Fen, Cambridge ( x 480). (diam. trich. 1'5 2'5/i fig. 155 B), and S. tenu-
;

issima Kiitz. (
= S. subsalsa CErsted) is often
abundant in salt and brackish water, rarely occurring in fresh water (diam.
trich. 12 u.
 RivulariacecB 337

Sub-order 2. TRICHOPHORE^E.
This is a small group containing some of the most character-
of the blue-green Alga3.
istic The trichomes are strongly attenu-
ated either towards one end, or from the middle towards both
ends, and in cases they are sheathed.
all The sheaths are generally
strong, rarely hyaline and delicate, often lamellose, and fimbriate
or ocreate along their margins. In certain genera, such as in
Dichothrix and Ammatoidea, a false branching is present.Hetero-
Asexual
cysts are present in some genera, but absent from others.
reproduction takes place by the formation of hormogones, but
in
Gloeotrichia spores are developed from the basal cells next the

heterocysts.
There are two sharply differentiated families :

Family 1. Rivulariacece. Trichomes attenuated from the base to

the apex, which is piliferous heterocysts basal (rarely absent).
;

Family 2. Camptotrichacece. Trichomes attenuated from the

middle towards each extremity ; heterocysts absent.

Family 1. RIVULARIACE^J.
Algae of this family are fairly abundant in mountainous dis-
tricts, and they also occur sparingly in the less elevated parts of
the country. They are found principally on the dripping rocks of
waterfalls, cataracts and streams, or at the margins of rocky lakes.
They form soft felt-like expanses, or hard hemispherical masses,
generally of a brown colour, but a few of them
occur as blue-green
nodules attached to the stems and leaves of submerged plants.
The trichomes are all attenuated to a hair-like point, and at
the broad base, in all except a few species, one or two heterocysts
are located. are present in
Occasionally intercalary heterocysts
addition to the basal ones. The sheath is tubular, gelatinous or
membranous, and is frequently strongly lamellose.
species In some
the lamella? of the sheath become dilated upwards, thus giving
the exterior of the filament a fimbriate appearance. It frequently
happens that the sheaths of adjacent filaments
become fused to
such an extent that their individuality is lost. The usual colour
of the sheaths is yellow or yellowish-brown, and in some cases
v

they are indurated with carbonate of lime.

The filaments exhibit a false branching due to the growth of
w. A.
338 Myxophycece
new trichomes from some of the inferior cells of the original

trichome, the new growth often occurring immediately above an

intercalary heterocyst. It is in Dichothrix that the false branching

most marked.
and fusion of the sheaths are
Asexual reproduction takes place by the formation of hormo-
of the trichome being lost.
gones, the hair-like apical portion
Asexual spores commonly arise in Gloeotrichia (and also in Calothrix)
in close apposition to the basal heterocysts.
There are five British freshwater genera, which can be distinguished as
follows :

A. Without heterocysts. Filaments free, very narrow,

forming a thin stratum Amphithrix.
B. With heterocysts.
* Filaments
free, simple or forming a dichotomous thallus.
t Filaments simple or falsely branched; branches
distinct and free Calothrix.
ft Filaments branched branches several (2
; 6)
within a common sheath Dichothri.r.
** Filaments
forming a hemispherical or globular thallus, closely
united by mucus.
t Filaments radiately disposed in a tough,
globose or hemispherical, attached thallus.
Spores unknown Rivularia.
tt Filaments radiately disposed in a soft, globose,
free-floating thallus. Spores regularly pro-
duced Gloeotrichia.

Genus Amphithrix Kiitz., 1843; em. Born. & Flah., 188C.

The filaments form a thin expanded stratum of a purple or violet
colour, which consists of two layers. The inferior layer is com-
posed of densely intricate filaments, or of minute radiately disposed
series of cells the superior layer consists of simple, erect filaments,
;

closely packed together and attenuated to fine points. The sheaths

are thin, close and continuous, and there are no heterocysts.
A. janthina (Mont.) Born. & Flah. occurs on wet rocks and is known from
W. Yorkshire; thickness of filaments 1'5 2'25 p.

Genus Calothrix [Mastigonema Schwa'be, 1837

Ag., 1824. ;

Mastigothrix Kiitz., 1843; Schizosiphon Kiitz., 1843 (in part);

Sympkyosiphon Kiitz., 1843 (in part).] The filaments are simple
or slightly branched, forming penicillate tufts or soft velvety

expansions, which are generally attached to submerged rocks and

stones. The heterocysts are basal or intercalary, and in a few
species they are absent. The plants often exhibit a slight
 Rivulariacece 33!)

branching, caused by the close application of the base of one

filament to the side of another, the sheaths being continuous.
The upper and more attenuated parts of the filaments are, how-
ever, always free. Borzi has 1
observed the formation of basal,
seriate spores in one species.
There are about six British freshwater species, C. parietina (Nag.) Thur.
being the most frequent (thickness of trichomes 6 12 p; fig. 156 and B). A
C. fusca (Kiitz.) Born. & Flah. and C. epiphytica W. & G. S. West are epi-

phytes on other larger Algpe, such as Vauchena or Batrachospermum.

Fig. 156. A and B, Calothrix parietiiici (N;i<, .) Thur., from Arncliffe, W. Yorks.
r

C, Dichothrix intfrntpta W. & G. S. West, from Slieve Donard, Ireland.

D, D. Orsiniiinii (Kiitz.) Born. & Flah., from Langdale, Westmoreland. (All
x420.)

Genus. Dichothrix Zanardini, 1858. In this genus the filaments

are moreor less -dichotomously branched, several trichomes with
their sheaths being enclosed within an outer common sheath. The
heterocysts are basal or intercalary, and in one species they are
absenf. The plants generally occur in penicillate tufts on dripping
rocks.
1
Borzi in Nuovo Giornale Botau. Ital. 1882, xiv, p. 374.

99.
340 Myxophycece
There are five British species, all of which are rare. They occur princi-
pally on dripping rocks in mountainous regions. D. Nordstedtii Born.
& Flah. and D. Orsiniana (Klitz.) Born. & Flah. (thickness of filaments
10 12 /u, of trichomes 6 7'5 p fig. 156 D) are widely distributed, and D.
;

W.
i & G. S. West is known from the Mourne Mts, Down, Ireland
interrupt
(fig.
156 C).

Genus Rivularia (Roth) Ag., 1824 (in part); em. Thuret, 1885.
[Zonatrichia J. Ag., 1842 ;

Limnactis Klitz., 1843 ;

Schizosiphon Kiitz., 1843

(in part).] The plants
consist of a globose or

hemispherical thallus, of a

tough character, composed

of radiating filaments which
'
are repeatedly branched.'
The thallus is attached to
submerged plants (such as
Chara, Myriophyllum, or
the bases of the stems of
Phragmites) or to the stones
of streams and cataracts,
and is sometimes indurated
with lime. The heterocysts
are basal and the extremi-
ties of the filaments are

piliferous. Spores have not

"** '"*''
been observed.
tt_-^e jgSjVj

There are four freshwater

A species occurring in Britain, of
Fig. 1T>7. C, Rivularia Biasolettiana
Menegh., from Aruclitfe, W. Yorks. A, nat. ;
which R. hematites (D. C.) Ag.
size, on surface of stone; B and C, x 480.
[= Zonatrichia calcarea (Eng.
D and E, R. mimHula (Kiitz.) Born. & Flah., Bot.) Endlicher] is the most
from Chippenham Fen, Cambridge; D, nat.
size, on stem of Phragmites; E, x 480.
frequent, sometimes occurring
in quantity attached to the
stones of mountain streams in limestone districts. R. dura Roth and R.
minulnlit (Kiitz.) Born. & Flah. (thickness of trichomes 2 12'5^; fig. 157 D
and E) are occasionally met with attached to submerged plants, forming small
globose masses of a blue-green or greenish-black colour.

Genus Glceotrichia J. Ag., 1842. The thallusis globose and

free-floating, solid when young, but inflated and hollow when

 Camptotrichacece 341

old. The filaments radiate from the centre outwards, and, as in

Rivularia, exhibit a false 'branching.' The sheaths are only
conspicuous near the base of the trichomes, being gelatinous and
confluent near the periphery of the thallus. The trichomes are
strongly attenuated from the base, and are more or less torulose.
From the cell immediately above the basal heterocyst elongated
cylindrical spores
are developed, which remain for some time
within the basal part of the sheath.
There are two British freshwater species, G. Pisum (Ag.) Thur. and
G. natans (Hedw.) Eabenh. The former possesses a thallus which does not
reach a greater diameter than 2 mm., but the latter is a larger species with
a thallus up to 10 cms. diameter. <!. I'lsum is one of the Algae
frequently
"
concerned in the phenomenon of the breaking of the meres."

Family 2. CAMPTOTRICHACEJE.
Thisa small family sharply
is marked off from the rest of the
Trichophoreae by the attenua-
tion of the filaments from the
middle towards each extremity.
The plants are epiphytic and
there are no heterocysts. The
sheath may be very thin and
delicate, or thickand lamellose.
Only two genera are known, one
of which (Camptothrix) is a genus
of small tropical epiphytes.

Genus Ammatoidea West &

G. S. West, 1897. The filaments
are epiphytic and exhibit a false
ramification comparable with that
met with in Rivularia. They are
suddenly bent in their median or
widest portion, the two more or
less parallel extremities being

gradually attenuated to piliferous.

apices. The sheaths are firm and
lamellose, in young filaments
colourless, but in older ones of a
Fig 158 ^mmatoidea Xnrm.niii
vpllovv brown colour
ve The tri- ' '

W. & G. S. West, from Dartmoor,

chomes exhibit a slight constric- Devonshire (
x 420).
342 Myxophycece
tionbetween their median cells, and the apical cells are about six
times longer than their diameter.
A. Normanii W. & G. S. West is known from Devonshire as an epiphyte
on Batrachospermum moniliforme. Thickness of filaments 5'5 12'5/i, of
trichomes 3'5 5'5 p fig. 158.
;

Order II. COCCOGONE^E.

In this order the plants are unicellular or colonial, commonly
occurring as colonies of unicells enveloped in a copious gelatinous
investment. The colonies vary much in size and shape, being
or
spherical, ellipsoidal, expanded gelatinous masses. The cells are
of varied form and are disposed in a variety of ways within the
mucous envelope. The latter is sometimes conspicuously lamellose,
but more often it is hyaline and structureless.
The Coccogoneas constitute the lowest group of the Myxophyceas,
and indeed they are the most primitive of all the Algse. They
occur free-floating, or more rarely as epiphytes, and some of them
form gelatinous masses on wet rocks. Not a few are regular and
abundant constituents of the freshwater plankton.
The normal method of multiplication is by simple cell-fission,
the larger colonies ultimately breaking up to form smaller ones.
Sometimes young colonies arise at the sides of the older ones by
a process of budding. Rounded asexual spores or gonidia have
been observed in some species, being formed within the wall of a
mother-cell.

Zoogonidia are unknown.

There are two well-marked families.

Family 1. Chamaisiphoniacece. Cells epiphytic, with a distinct

base and apex ; reproduction only by the formation of gonidia.
Family 2. Chroococcacece. Cells or colonies free-floating or form-

ing a gelatinous stratum, very rarely epiphytic, not differentiated into

base and apex ; multiplication by simple cell-division (very rarely by
gonidia).

Family 1. CHAM.SISIPHONIACE^E.
The plants of this family are epiphytes, generally occurring in
clusters around the filaments of larger Alga3. They are in all
cases differentiated so as to present a distinction between base
and apex, and reproduction occurs by the formation of a number
 Chroococcacece 343

of non-motile spores or gonidia from the contents of a mother-cell.

Most Algae of this family are marine, the only British freshwater
genus being Ghamcssiphon.
Genus Chamsesiphon A. Br. & Grim., 1864. [Indus. Sphcero-
gonium Rostaf, 1883.] The cells are small, ovoidal, pyriform, or
with very thin cell-walls. The cell-contents are homo-
cylindrical,
geneous and of a blue-green, violet, or yellow colour. The cells
are attached by their base and generally widen upwards to their
free apex. The gonidia
are successively cut off from the upper

part of the which has become a gonidangium, gradually

cell

escaping from the open apex.

Fig. 159. Ckamce siphon incntstans Grim., on a filament of Rhizocloniuiit,

from Heaton, W. Yorks. ( x 416).
%

Ch. confervicola A. Br. found as an epiphyte on Chcetomorpha, Rhizoclo-

is

nium, Vaucheria, etc., and Ch. incrustans Grun. (diam. of cells 3'5 4'8/i;
fig. 159) often thickly covers filaments of Rhizoclonium and CEdogonium.

Family 2. CHROOCOCCACE^.
This the largest family of the Coccogonese, and includes a
is

great variety of unicellular and colonial blue-green Alga?. They

are abundant in all kinds of damp and wet situations, frequently

forming a gelatinous stratum on the damp surfaces of dripping

rocks. The cells
vary much
in shape in the different genera, and
the colonies assume different forms according to the direction of
division of the cells. In some the cells divide in every direction
of space within a gelatinous envelope, producing an irregular

colony, often of large size. In others the cells divide only in two
directions in the same plane, giving rise to a tabular colony; and
in others cell-division takes place in one direction only. In one
genus (Tetrapedia) the cells are flattened and they possess a
striking symmetry of form. Synechococcus and Tetrapedia are the
only genera which are commonly destitute of a mucous envelope.
In all others the cells are invariably embedded in a mass of mucus,
344 MyxophycecB
which varies much in its consistency. It may be firm and lamellose,
or very hyaline and diffluent. The cells often contain red, orange,
or violet pigments.

Reproduction has been observed in a few species to take place

by the formation of non-motile spores or gonidia within a goni-
dangium.
There are two sub-families :

Sub-family I.
Chroocystece. Epiphytes on larger Algse, with a
well-defined dorsiventrality.

Sub-family II. Chroococceee. Free-floating or forming a gelatinous

stratum, with no dorsiventrality.

Sub-family I. CHROOCYSTE^E.
This sub-family includes only a single genus, the plants of
which differ from the rest of the Chroococcacea3 in being epiphytic,
and in the possession of distinct upper and lower surfaces. They
are likewise characterized by the possession of prominent bristles.
Genus Gloeochgete Lagerh., 1883. [Schrammia Dangeard,
1889.] The cells are globose or sub-globose, enveloped in a wide

Fig. 160. Glaochcete Wittrockiana Lagerh. A, from Cray Moss, W. Yorks.;

B, from Pilmoor, N. Yorks. ( x 416).

mucous coat, and they occur singly, or in twos or fours, attached

to larger filamentous
Alga3. More rarely they are attached to the
 Chroococeacew 345

leaves and stems of submerged Mosses or Phanerogams. Each

cell is furnished at upper pole with one or two long, thin
its

bristles, Avhich sometimes give off small branches or spurs near

their apices. The primitive chromatophore is bell-shaped, of a
brilliant blue-green colour, and there is a clear space in the centre
:
of the cell. Lagerheimin his original description of the genus
"
says nucleus singulus." The mother-cells give rise to two or
four daughter-cells on division.

Many authors have placed this Alga in the Chlorophyceae, or

have recorded a chlorophyceous Alga under the name of Glceochcete.' '

The genus as I have often found it, and as here figured, most
certainly belongs to the Chroococcacese.
(.t'L Wittrockiana Lagerh. is known from several parts of the British
Islands, occurring as an epiphyte on Vaucheria, Cladophora, or the leaves of
Sphagnum. Diam. of cells 6 21 p length of bristles 96 260 fj. fig. 160.
; ;

GL bicornis Kirchn. has a pair of bristles attached to each cell.

Sub-family II. CHROOCOCCE^.

The sub-family Chroococcese embraces almost all the unicellular
and colonial Myxophycea?. They are unquestionably of a lower type
than any other of the Alga3, and some of them bear resemblance
to certain of the Bacteria. The cells display considerable variety
of form, and with the exception of the genera Synechococcus,

Tetrapedia, and to a certain extent Merismopedia, they are

enveloped in a copious covering of mucus. The mucous coat is
sometimes firm and lamellose, as in some species of Chroococcus
and Glceocapsa, but more often it is an ample, homogeneous covering,
very hyaline in character. It often happens that the coverings
of numerous have fused together, thus forming gelatinous
cells
colonies of various sizes. Some of these colonies are large macro-
scopic masses, containing many thousands of cells, and possessing
a tough exterior. In some genera, such as Merismopedia and
Gcelosphcerium, the colonies are of limited size and of definite
shape.
Multiplication takes place by the repeated division of the cells
and the final fragmentation of the original colony, each fragment

growing in size and repeating the processes. In the spherical

colonies of Cvelosphcerium a kind of budding takes place by means
1
Lagerheim in Nuova Notarisia, 1890, p. 231.
346 Myjcophycece

of which a developed from the side of the old one,

new colony is

ultimately becoming separated

from it. Cell-division may be in
one direction only, in two directions in one plane, or in every
direction of space.

Reproduction occurs in Gomphoaphceria by non-motile spores

1
or gonidia .

There are thirteen British freshwater genera, which can be arranged as

follows :

A. Cell-division in one direction only.

* Cell-division transverse.
t Cells enveloped in a wide mucous coat.
\ Cells elongated, each
with a mucous coat Gloeothece.

\\ Cells little longer than broad, enveloped

in a common mucous investment ... Aphanothece.
t 1 Cells destitute of mucus ........................ Synechococcus.
** Cell-division cells enveloped in mucus Dactylococcopsis.
oblique ;

B. two directions in one plane.

Cell-division in
* Cells colonies
globular or rounded-quadrate ;

often large ............................................. Merismopedia.

** Cells of a definite
symmetrical form, solitary,
or forming small colonies ........................ Tetrapedia.
C. Cell-division in all directions of space ;
cells enveloped in mucus.
* Cells
forming large colonies.
t Cells arranged at or towards the periphery of spherical
colonies.

J Cells closely
and regularly arranged ... Codosph<xrium.
\\ Cells geminate and sparsely scattered,
markedly pyriform in shape ............ Gomphosphcena.
tt Cells densely aggregated in globose, elon-
gated, or clathrate colonies ............... Microcystis.
ttt Cells aggregated to form irregular gelatinous colonies.
j Individual mucous coats clearly evident
round each cell ........................... Glceocapsa.
\\ Cells enveloped in a common mucous
covering .................................... Aphanocapsa.
JJ| Cells arranged in a compact gelatinous
stratum .................................... Porphyridium.
** Cells more or less
solitary, or forming very
small colonies ....................................... Ckroococcus.

Genus Gloeothece Nag., 1849. The cells are cylindrical and

a thick mucous coat,
elongated, each one being surrounded by
which sometimes shows indications of lamellation. Cell-division

1
Schmidle in Berichte Deutsch. Botan. Gesellsch. 1901, xix.
 Chroococcacece 347

is
only in one direction, and a number of cells are usually associ-
ated to form a small colony.

Species of this genus occur on wet rocks, among wet mosses, and in bog-
pools. Gl. Nag. (length of cells 1O5 18 p.; breadth 1'3 2-3 /i;
liiit'firis

fig. 161 A) and Gl. confluens Nag. (length of cells 5-7 7'5 p. breadth 2-63 p.; ;

fig. 161 B) are the

most frequent British species.
lihabdoderma lineare Schmidle 1 seems very closely allied to Gloeothece
linearis Nag., if not actually identical with it.

Genus Aphanothece Nag., 1849. This genus only differs

from Gloeothece in the ag-
gregation of large numbers
of cells within a common
mass of mucus. The cells

are cylindrical and longer **<&

than their diameter.
A. microscopica Nag. (length
of cells /z,
58
breadth 3'5
4/x ; fig. 161 C) and A. saxicola
Nag. are the most frequent
species. They are found in
bog-pools, at the margins of
lakes, and on wet rocks.

Genus Synechococcus
Nag., 1849. The cells,
which are cylindrical with
hemispherical apices, are
larger than in the preced- Fig. 161. A, Gloeothece linearis Niig., from
Old Cote Moor, W. Yorks. B, Gl. conjluens
ing genera, and are desti-
Nag., from near Settle, W. Yorks. C, Aphano-
tute of the outer mucous thece microscopica Nag., from Withiel, Corn-
coat. occur free-
wall. D and E, Synechococcus major Schroet.,
They from Adel Bog, W. Yorks. (All x 450.)
floating in ponds, ditches,
and bog-pools, often in considerable quantity. The cell-contents
are usually of a brilliant blue-green, rarely of a rose-purple colour,
and contain numerous large granules.
S. ceruginosusNag. and >$'. major Schroeter (length of cells 26 29 p. ;

breadth 15'5 17'5//; fig. 161 D

and E) are the most abundant British
species, the latter often occurring in quantity in bogs.

Genus Dactylococcopsis Hansg., 1888. The cells are generally

associated to form small colonies, rarely solitary, and in many
1
Schmidle in Berichte Deutsch. Botan. Gesellsch. 1900, xviii, p. 149, t. vi,
f. 811.
348 Myxophycece
cases the colonies are embedded in a copious mucus. The cells
are elongated, fusiform or lanceolate, with attenuated extremities,
or sometimes sigmoidal in form. The chromatophore is almost
homogeneous, of a pale blue-green colour, and somewhat lateral in

position. Division of the cells occurs much as in Dactylococcus,

by oblique septation.
D. montana W. & G. S. West (length of cells 8-611-5 M breadth 3'5
,
4/*;
fig. 162 A) occurs in bog-pools amongst Sphagnum. D. rliaphidioides Hansg.
is known from the plankton of Lough .Neagh.

Genus Merismopedia Meyen, 1839. This is one of the most

striking genera of the
Chroococcacese, consist-
)t a
A rectangular
colony, the cells of which

^ 'Y>
are arranged in rectilinear
series. Cell-division

place in two directions

and the cells appear to
be usually arranged in
takes

groups of four. The cells

are globose, ellipsoidal, or
338388
oblong, sometimesslightly
Fig. 162. A, moiitumi
Dactylococcopsis angular by compression,
W. & G. S. West, from Widdale Fell, W. Yorks. and the cell-contents are
B, Merismopedia glauca (Ehrenb.) Nag., from
Thursley Common, Surrey. C, M. elegans homogeneous.
A. Br., part of a large colony from Wicken
Fen, Cambridge. D, Tetrapedia Eeinscliiana J/.glauca (Ehrenb.) Nag.
Arch., from near Goring, Oxford. (All x 450.) is the commonest British
species, occurring in ponds,
and lakes; diarn. of cells 3'3 3'8/x;
ditches, bogs fig. 162 B. M. punctata
Meyen and M. ceruginea Breb. are not infrequent in stagnant water. M.
elegans A. Br. the largest species of the genus and is known from few
is

localities in the British Islands; diam. of cells 6'5 9'5/x; fig. 162 C. The
colonies of M. glauca and M. elegans often reach a large size (diam.
up to
220 fj.)
and may contain as many as 1856 cells.

Genus Tetrapedia Reinsch, 1867. In this genus the cells are

ofsome definite and symmetrical shape, often constricted into two
equal half-cells. The cell-wall is firm and the cell-contents are
homogeneous. In some species the cells are almost invariably
solitary, but in others they are grouped in flat colonies similar to
those of Merismopedia.
 Chroococcacece 349
All the species are rare, but T. Reinschiana Arch. (diam. of cells 5*5 6 p.;
fig. 162 D) and T. glaucescens (Wittr.) Boldt occur in lakes and bo^- pools.
T. setigera Arch, is a beautiful little species known from X. W. Scotland and
W. Ireland, and which has been erroneously referred by some authors to the
chlorophyceous genus Tetraedron. The cell-contents of Tetrapedia s<.'ti;f<-i->r
are homogeneous and of a pale blue-green colour.

Genus Coelosphaerium Nag., 1849. The cells are more or less

closely grouped form a hollow, spherical colony. They are em-

to
bedded in a mass of mucus and are arranged just within the
periphery of the spherical mass. In form they are globose, ellip-
soidal, or ovoidal, and the cell-contents are granular, with so-called

gas-vacuoles.
The genus is a common constituent of the freshwater plankton, and is

also found frequently in large ponds. C. Kutzingianum Nag. (diam. of colony

48 90/ij diam. of cells 3 3'8 /*; fig. 163 A) is the most abundant species,
although C. Niigelianum linger and C. minutissimum Lemm. both occur in
quantity in the British freshwater plankton.

^"A-
-o,
oo9

Fig. 163. A, pcelospJusrium Kiitzingianum Nag., from the plankton of Lough

Neagh, Ireland. B, Gomphosphieria aponina Kiitz., from Kestou Common,
Kent (x350).

Genus Gomphosphaeria Kiitz., 1836. In this genus the

colonies contain fewerand more scattered cells, which are disposed
chiefly towards the periphery of a globular or ellipsoid mass of
mucus. The cells are grouped in pairs and are distinctly pyriform
'
in shape. Schmidle has observed the formation of microgonidia.'
The entire colony is solid and the cells divide alternately in three
directions.

G. aponina Kiitz. (diam. of colonies 64 78 p; length of cells 8'">

11'5/u;
fig. 163 B) is not uncommon in ponds, lakes, and stagnant ditches.

Genus Microcystis Kiitz., 1833. [Polycystis Kiitz., 1845;

Clatkrocystis Henfrey, 1856.] The cells are small, mostly globose,
350 Myxophycece
and are densely aggregated to form solid colonies of variable
shape. In some species the colonies are globose, ellipsoidal, or
oblong ;
in others they are
much elongated and in
;

others they become cla-

thrate and almost anasto-
mosing. The cell-contents
are blue-green, olive-green,
or rose-purple in colour,
and often contain
B vacuoles.

Fig. 164. A, Micrwystis stagnalis Lemrn.,

M. marginata Menegh.
from the plankton of Lough Neagh, Ireland. (diani. of cells
2'6 2'8/i; fig.
B, If. marrjinata Menegh., from Old Cote 164 B), M. Flos-aquas (Wittr.)
Moor, W. Yorks. ( x 450).
Kirchn., M. elabens (Breb.)
of cells 11-5 /* tig. 164 A) are all more
Kiitz., and J/. stagnalis Lemm. (diam.
;

or less common in bogs and lakes. M. roseo-persicinus (Kiitz.) often occurs

in quantity in ponds and ditches which contain much decaying vegetation.
M. =
ceruginosa (Kiitz.) [ Clathrocysti* ceruginosa (Kiitz.) Henfrey]
often occurs

in prodigious quantity in ponds and in the plankton of lakes.

The three genera Microcystis, Polycystis and Clathrocystis are not suffi-
The differences between them
ciently distinct to warrant
their separation.
are only differences of degree.

Genus Gloeocapsa Kiitz., 1843 em. Nag. 1849. The cells are
;

with a thick integument, which is frequently

globose and furnished
lamellose. The daughter-cells which arise by the division of the
mother-cells are generally retained as part of the colony, a larger
of the cells. In
integument surrounding the individual envelopes
this manner the colonies become of large size and frequently form
a gelatinous stratum. The integuments of the cells are sometimes
or red.
colourless, but they may be yellow, brown, blue, violet,
Cell-division takes place in all directions and the colonies are
most ThTct-walled have been observed
resting-spores
irregular.
in some species.
Kuntze and, following him, certain other authors, have identi-

fied Gloeocapsa Kiitz. with Bichatia Turp. (1827), but to my mind

this identification is uncertain.

There are about 20 British species, some of which are of doubtful specific
2/*) is the
smallest
distinctness. Gl. punctata, Nag. (diam. of cells 1-5
species. (11.
poli/d<'nnatica Kiitz. is remarkable for the larnellation of the
integuments (fig. 165 C E). Gl. Magma (Breb.) Kiitz. with golden-yellow or
 Chroococcacecf 351

In-own integuments (fig. 165 B), and Gl. Ralfsiana (Mass.) Kiitz. with bright
red or purple integuments, are two of the best-defined species. Most of the
-species occur on wet or dripping rocks, generally in association with other
Myxophycese.

Genus Aphanocapsa Nag., 1849. The cells are globose and

aggregated toform small
colonies within a com-
mon homogeneous in- {
9
tegument. The genus
only differs from Apha-
nothece in its globose
00
cells. The integument
p B
may be colourless or
CG
tinted brown or blue- _Q
green, and the cells are
of a blue-green or olive-

green colour.
There are four British
species occurring both in
stagnant water and on wet
rocks. A. Grevillei (Hass.)
D
Rabenh. is the most fre-

quent diam. of cells 3 '4 165. A, Aphanocapsa Grevillei (Hass.)

;
Fig.
4-5 /A; fig.165 A. Eabenh., from Helln Pot, W. Yorks. B, Gltxo-
cap.srt M/III ma (Breb.) Kiitz., from Boston Spa,

Genus W. Yorks. C E, Gl. polyde.rmatica Kiitz., from

Porphyri- Boston Spa, W. Yorks. (All x 450. )
dium Nag., 1849. The
cells, which are closely arranged to form a thin gelatinous stratum,
are globose or angular by compression. The stratum consists of
many layers of cells, and the cell-contents are of a reddish-purple
colour. Cell-division takes place in all directions.

P. cruentum (Ag.) Nag. is a common Alga, forming a thin slimy stratum

of a dark red colour on damp ground and near the base of damp walls. The
cells are 7 9 p.
in diameter.
This Alga was at one time placed in the Chlorophyceae and has since been
relegated to the Khodophycese. I agree with Hansgirg, however, in thinking

it much better placed in the Myxophyceae. There are many of the Myxo-
phyceae which possess as much red or purple pigment as Porphyridiwrn, and
moreover, the latter genus is generally found in association with blue-green
Algae. It is more nearly allied to Aphanocapsa than any other genus of

Algae.
352 Myxophycece
Genus Chroococcus Nag., 1849. In this genus the cells are
globose or more or less angular, solitary or associated in simple
families. They are free-
floating or mixed with
other blue-green Alga? to
form a stratum on wet
rocks. The integuments
are firm and often wide,

homogeneous or lamel-
lose, generally colourless,
but sometimes of a yellow-
ish-brown tint. The cell-
contents are granulose, of
a brilliant blue-green
colour, or more rarely
violet, olive-green, or
Fig. 166. A, Chroococcus giganteus West,
yellow-green.
from Bowness, Westmoreland. B, Ch. turgidus
(Kiitz.) Nag., from Slieve Donard, Down, Ireland. Ch. turgidus (Kiitz.) Nag.
C and D, Ch. schizodermaticus West, from near
isthe most widely distributed
Windermere, Westmoreland. (All x 450.)
species, often occurring in
quantity in Sphagnum-bogs ;
diam. of cells 13 25 p; fig. 166 B. Ch. cohcerens

(Breb.) Nag., Ch. giganteus West (fig. 166 A), Ch. minor (Kiitz.) Nag. and
Ch. pallidus Nag. are not infrequent in ponds, lakes, and bog-pools. Ch.
schizodermaticus West (fig. 166 C and D) is remarkable for its tough lamellose
integuments, the layers of which are gradually split off and shed. Ch. lim-
neticus Lernm. is confined to the freshwater plankton.
 INDEX.
[Synonyms are printed in italics, and the numbers in strong type
refer to the descriptions of the genera, families, orders, etc.]

Acanthococcus Lagerb., 203. Amphora Ehrenb., 298, 299.

Achnanthaceas, 289, 290. ovalis Kiitz., 269, 299 (fig. 141
Achnauthes Bory, 263, 289. B and C), 300.
coarctata Breb., 290. ovalis var. pediculus Kiitz., 300.
exilis Kiitz., 290. Anabasna Bory., 311, 312, 314, 317, 325,
flexella (Kiitz.) Breb., 269 (fig. 327.
125 B), 290. circinalis Eabenh., 328.
Hungarica Grun., 2110 (fig. 135 Flos-aquae Breb., 328.
A C). inaequalis (Kiitz.) Born. & Flak.,
linearis W. Sm.,
290. 328 150 A D).
(fig.
loiigipes C. Ag., 268. oscillarioides Bory, 328.
microcephala Kiitz., 290. Ancylonema Bcrygr., 144, 149.
Achnanthidium Kiitz. (sect, of Achnan- Nordenskioldii Berggr., 5, 51.
thes), 289, 290. Androgonidia, 191.
Acbnanthoideae, 280, 289291. Androsporangium, 61.
Actinastrum Lar/crli., 218, 224. Androspore, 61.
Hantzscbii Layerli., 225. Ankistrodesmns Corda, 159, 218, 221,
Ajuga,.199. 222225.
Akinetes, 15. Arch., 222, 223.
<ici(tixtiimiit>

Akonta, 32. biplex (Eeinsch) nob., 224.

Alocasia, 55. couvolutus (Pabenh.) nob., 221.
Alternation of generations, 18. falcatns (Corda) Half's, 53, 223 (fig.
Amblystegium exannulatum, 4. 94 A),
falcatum, 4. falcatus var. acicularis (.-1. Jlr.)
glaucum, 4. nob., 223 (fig. 94 B and C).
scorpioides, 4. falcatus var. duplex (Kiitz.) nob.,
Ammatoidea W. a; <1. S. West, 38, 337, 223.
341. falcatus var. mirabilis nob ,
223
Normanii W. & G. S. West, 341 (fig. 94 E), 224.
(tig. 158), 342. falcatus var. spiralis (Turn.) nub.,
Amceba, 4, 145, 230. 224.
Amphk-ampa Ehrenb. (1849), 288. falcatus var. spirilliformls nol>., 22-1.
Ainphicampa Rabeiih. (18(54), 296. falcatus var. tumidus nob. 223 ifitr. ,

Arupbipleura Kiitz., 263, 292, 295. Ill

D), 224.
pellucida Kiitz., 295 (fig. 137 C). /n.-ii/iirinia Corda, 222, 223.
Ainpbiprora Khrcnb.. 291, 292, 296. Pfitzeri (Sch rH,], '/) nob.. 223 (fa.
ornata Bail., 296. 94 G
and H), 224.
paludosa W. Sm., 296 (fig. 138 setigerus (Schroder) >iol>., 223 (fig.
B and C). 94 F), 224.
Ampbithrix Kiits., 338. Anodonta, 4.
jantbina (Mont.) Born. <&Flah.,338. Anorthoncis Grun., 290.

W. A. 23
354 Index
Antheridium, 17. Bacillariese, 5, 6, 8, 11, 15, 16, 31, 32,
Antherozoids (or spermatozoids), 17. 260305.
Antlioceros, 325. Bacteria, 3, 316, 332, 345.
Aphanizomenon Morren, 308, 311, 326, Bangia, 36.
328. Bangiaceae, 35, 98.
Flos-aquas (L.) Ealfs, 328. Batrachospermurn Roth, 18, 20, 36, 38,
Aphanocapsa Nfig., 346, 351. 39, 339.
Grevillei (Hass.) Rabenh., 351 (fig. atrum (Dillw.) Harv., 38
165 A). Boryanum, 35.
Aphaiiocluete A. Br. Berth.
; Huber, ; niom'liforme Roth, 37 (fig. 1 A),
71, 72, 89, 182. 38, 342.
globosa var. minor Haiisg. 182. , vagum (Roth) Aq., 37 (fig. 1 B and
pilosiasiina Schmidle, 72. C), 38.
repens A. Br., 72. Beuecke, 264.
Aphauothece Nfig., 13, 346, 347, 351. Bennett, 1.

microscopica Niig., 347 (tig. 161 C). Bennett & Murray, 111, 125.
saxicola Nfig., 347. Berggren, 5.
Apiocystis Nag., 51, 239, 244. Bessey, 148.
Brauriiana Nag., 244 (tig. 112). Beyerinck, 230.
Aplanogametes, 16. Bichatia Turp. 350. ,

Aplanospores, 15. Biddulphoidese, 273.

Aptogonum Ealfs, 177. Binuclearia Wittr., 75, 80.
ArchegoniatsB, 30. tatraua Wittr., 80 (fig. 25), 81.
Archer, 1, 141, 142, 223. Blackmail, 2, 21, '22, 24, 27.
Archiplast, 317. Blackmail & Tansley, 32, 33, 102, 188,
Archiplastidese, 309, 316, 317352. 248.
Areschoug, 75. Blasia, 4.
Arisarum, 55. Boergesen, 1, 99, 143.
vulgare, 54. Bohlin, 21, 2629, 32, 100, 108, 248,
Arthrodesmus Ehrenb., 137, 144, 151, 253, 254, 256.
169, 170. Boldt, 143.
bitidus Breb. var. truncatus West, Borge, 1, 12, 125, 147.
170 (fig. 64 H J). Bornet, 1.
convergens Ehrenb., 171. Borzi, 2, 21, 28, 48, 78, 92, 182, 199,
Incus (Breb.) Hass., 170 (fig. 64 24*, 311, 314, 326, 339.
A C), 171. Botrydiaceffi, 29, 30, 249, 258, 259.
Incus W. & G. S. West,
var. Ealfsii Botrydiua Breb., 247.
170 (fig. 64 D). vulgaris Breb., 247.
Incus var. validus W. d: G. S. West, Botrydiopsis Borzi, 28, 29, 254.
170 (fig. 64 E). Botrydium Wall, 12, 18, 28, 29, 51, 258.
octocornis Ehrenb., 170 (fig. 64 granulatum(L.) Grev., 259 (fig. 122).
F aud G), 171. Botryococcus Kiitz., 235, 237.
Arthrodin Bafinesque, 159. Brauuii Kiitz., 237 (fig. 106), 242.
Arthrodieas, 148. calcareus West, 237.
Arthrosiphon Kiitz., 323. sudeticus Leiiini., 237, 242.
Arthrospira Stizenb., 315, 330, 332, 333, Boubier, 114.
336. Brady pus (Three-toed Sloth), 55.
Jenneri (Hass.) Stizenb., 336. Brand, 93, 104, 106.
Asexual reproduction, 14, 15. Brandt, 5, 311313.
Askenasy, 210. Breaking of the Meres, 315, 341.
Asterionella Hass., 287. Brewer, 6, 307.
fonnosa Hass., 2*7 (fig. 133). Bulbocha-te Ag., 5'2, 57, 58^ 65.
gracillima Heib., 287. gigantea Priniiah., 65.
Autocolouy, 25, 212. nana Wittr., 64 (tig. 15 C), 65.
Autospores, 14, 25, 212. Nordstedtii Wittr., 64 (fig. 15 B).
Auxospores, 268, 269. subintermedia A7/'r., 64 (tig. 15 A).
Azolla, 4, 325. Bumilleria Borzi, 29,' 253, 258.
pumila W. <(' G. S. West, 257
Bacilhiria (jnu'lin, 301, 302. (tig. 121 J), 258.
paradoxa Gmelin, 302. Biitschli, 266.
 Index 355

Callitriche, 205. Cb.BBtopb.ora pisiformis (Roth) Ag., 85.

Calocylindrus (Nag.) Kirclm., 166, 167. tuberculosa (Roth) ./</., 85.
Calothrix AH., 38, 312, 338. Chaetophoracese, 19, 26, :-io, 52, 60, 67,
epiphytica W. & (i. S. West, 339. 70, 8389, 93, 179.
fusca (Kiitz.) Born. & Flak., 339. Chsetophorales, 11, 15, 16, 19, 25, 30,
parietiua (AY/Y/.) T//MJ-., 339 (fig. 50, 54, 56, 6695, 98, 180. '
15G A and B). Chsetospbseridium Kl,'bnlui,52, 180, 181,
Campbell, 21. 182.
Camptothrix II'. G. S. West. 341.
<
globosum (Xordst.) Kleb., 182 (fig.
Camptotrichaceae, 337, 341, 342. 70 A and B), 183.
Campylodiscus Ehrenb., 3U3, 305. globosum var. depressant If. <&
Echineis Ehrenb., 305. G. S. W^t, 182 (fig. 70 C).
Hibernicus Ehrenb., 305 (fig. 145 minus Hansg., 182.
D and E). Pringsheimii Klebahn, 182.
Capsulococcus Bennett, 246. Chamassiphon A. Jir. <( Grim., 343.
crateriformis Jieitiiett, 246. confervicola, A. Br., 343.
Carotin, 308. incrustans Grim., 343 (fig. 159).
Carpogamous heterogamy, 16. Charnaisiphoniaceaa, 313, 342.
Carpogonium, 16, 17, 34. Chantransia Fries, 20, 38, 39, 42.
Carpospores, 16. corymbifera Thin:, 39.
Carteria Diesini/, 187. pygniffia Kiitz., 39 (tig. 2 A C).
multifilis (Fresen.) Dill, 187, 188 Scotica Kiitz., 39 (fig. 2 D).
(fig. A G). 73 Cbara, 340.
Castracane, 269, 270, 272. Characies, 30, 179, 199200.
Caulerpa, 25, 109. Characiopsis Borzi, 28, 29, 250, 251.
Cellulose, 51. minuta (A. Br.) Borzi, 251 (fig.
Central body (of Myxophycese), 309, 117 A).
310.' turgida W. & G. N. West, 251 (fig.
Centric, 273 279. 117 B Dj.
Centrosphagra Borzi, 199. Characiuni A. Br., 197, 200, 219, 251.
Facciolase Borzi, 198 (fig. 79 arnbiguum A. Br., 200.
B and D), 199. ensitbrme Herm., 200 (fig. 80 D).
Cephaleuros, 4, 13. ornithocepbalum A. Br., 200.
Cerasterias Eeinsch, 231. Pringsbeimii A. Br., 200 (fig. 80 A
longispina (Perty) W. & G. S. West, and B).
232. Sieboldii A. Br., 200.
rhaphidioides Reinsch, 232. subulatum A. Br., 200 (fig. 80 C).
Ceratodus, 7. Chlamydococcus A. Br., 189.
Ceratoneis Ehrenb., 288. pluvialis (Flot.) A. Br., 189.
Arcus (Ehrenb.) Kiitz., 288 (fig. Chlamydomonadese, 186 189.
134 D). Chlamydomonas Ehrenb., 3, 22, 23, 50,
Arcus var. Amphioxys (Rabenh.). 51, 187, 189, 193, 202.
De Toni, 2ss. Debaryana <;<ir*eh., 188 (fig. 73
Cerciditun elongatum Dang., 189. H and I).
Chaetomorpha Kiitz., 102, 103, 343. Ehrenlienjii Goroscb., 188.
sutoria (Berk.) Rabenh., 103 (fig. hyaliua, 23.
38). Kleinii Schmidle, 188 (fig. 73 J
Chffitonella Schmidle, 106. and K).
Goetzei Schmidle, 106. pulvisculus Ehrenb., 188.
Chaetopeltis Berth., 180, 181. Chloramceba Bohlin, 29, 30.
orbicularis Berth., 181. Chlorella Beyerincl;, 4, 22(5, 230.
Chaetopeltideae, 30, 52, 179, 180184. vulgaris Beyerinek, 230.
Chaetophora Schnnik., 19, 52, 67, 84, 85. Cblorobotrys Bohlin, 29, 249. 253.
ntli-fireti Tilden, 85. regularis (Wet) Hnhlin, 254 (fig.
Cornii Dunne. (Rotb) Ag., 85. 119).
elegans (Roth) Ay., 84 (fig. 27 C), Chlorocbytrium Colin, 197, 198.
85. .
Knyauum N: ijnm iml;i, I'.is.
1'itdii'if folia Ag., 85. Lemnse Cohit, 198 (fig. 79 A).
incrassata (Hudson) Huzen., 84 Chlorococcum Fries, 202, 245.
(fig. 27 A and B), 85. (Kiitz.) Grun., 246.

232
356 Index
Chlorococcum inftixionum (Schrauk) Cladophora flavescens Ay. ,
105.
Menegh., 246. i'racta Kiitz. ,
106.
105
regularc West, 254. glomerata (L.) Kiitz., (fig. 40),
Chlorogonium Ehrenb., 23, 187, 188. 106.
euchlorum Ehrenb., 189. Cladopboracese, 26, 30, 33, 50, 102
Chloromonadina (or Chlorornonadales), 106, 107, 252.
29, 248, 249, 253. Cladophorales, 11, 16, 26, 27, 30, 56,
Chloropbyceffi, 10, 15, 18, 19, 21, 22, 101108.
2630, 32, 33, 50247, 248, Cladothrix, 335.
270, 345, 351. Clathrocystis Henfrey, 349, 350.
Chloroplastids (or Chloroplasts), 12, 52. ceruginosa (Kiitz.) Heufrey, 350.
Chlorosaccus Luther, 29, 30. Cleve, 293.
Chlorosphccra Henfrey (1858), 229. Climacidium Ehrenb., 288.
Chlorosplucra Klebs (1883), 202. Climacosphenia Ehrenb., 283.
Chlorotheciaceffi, 29, 30, 249, 250252. Clonothrix nracillima W. & G. S. West,
Chlorothecium Borzi, 28, 29. 335.
Chlor-zinc-iodine, 51. Closteriese, 144, 150, 158.
Choaspis S. F. Gray, 134. Closteriopsis Lemm., 218, 224.
stictica (E-iuj. Bot.) 0. K., 134 longissima Lenun., 224.
(fig. 50)', 135. Closteriuin Nitzsch., 51, 136, 137, 140,
Chodat, 1, 2, 5, 21, 22, 26, 49. 90, 98, 144, 150, 158, 159, 162.
99, 195, 201, 216, 225, 228 acerosum (Schraitk) Ehrenb., 160
230, 232, 233, 242, 308, 311. (fig. 56 A), 161.
Chodat & Boubier, 50. acieulare T. West, 161.
Chodat & Cretier, 237. acutum Brt'b., 160 (fig.
56 E), 161.
Chodat & Grintzesco, 9. calosporum Wittr., 161.
Chodatella Lemm., 25, 232, 234. Ehrenbergii Meneijh., 139 (fig. 51 A),
breviseta W. <( <!. S. West, 234 161.
(fig. 102 F and G). gracile Breb., 161.
ciliata Lafjerh. var. araphitricha Kiitzingii Breb., 161.
103 Leibleiuii A'^., 139 51 B),
(Lagerh.) Chud., 234 (fig. (fig.
H and I),
161.
radiums (West) Lemm., 234. lineatum Ehrenb., 141 (fig. 52 F),
Choristocarpaceae, 45. 142.
Chromatophores, 12. moniliferum (Hurt/) Ehrenb., 161.
Chromoplastids (or Chromoplasts), 12. parvulum AV///., 160 (fig. 56 F), 161.
Chroinuliua Cieitk., 31. pronum lln'b., 161.
Cbroococcaceee, 2, 14, 314,342, 343 352. pusillum Hantzsch var. inono-
Chroococceae, 344, 345 352. lithuin Wittr., 161.
Chroococcus N<<<I., 345, 346, 352. Ealfsii Bre-b. var. hybridum llnbenh. ,

cobserens (Breb.) Nag., 352. 142.

giganteus West, 352 (fig. 166 A). rostratum Ehrenb., 161.
limneticus Lemm., 352. rostratum var. brevirostratum Went,
mucrticdcctis Ilabenh., 206. 160 (fig. 56 G).
minor (Kiitz.) AVV//., 352. striolatum Elirenli., 160 (fig. 56
pallidus Niiij., 352.
B D), 161.
schizoderrnaticus West, 352 (fig.
subtile Breb., 223.
16(5 C and D). turgidum Ehrenb., 161.
turgidus (Kiitz.) Nfig., 352 (tig.
Venus Kiitz., 161.
166 B). Cocci (of Myxophycese), 313.
Chroocystero, 344, 345. Coccogoneie, 318, 342352. %
Chnwlejins Ag., 95. Cuccomu.vu Schmidle, 2is.
clnuomonas Han*y., 32. Cocconeidacea>, 289, 290, 291.
Cliroothece Hdiisr).. 317. Coccoueis Ehrenb., 103. 261, 263, 290.
Richterianum Ilunxi/., 317. Pediculus Ehrenb., 291.
Chrysomonadinaceae, 31, 45, 46. Placeutula Ehrenb., 290 (fig. 135
Cienkowski, 29, 77, 82, 84. D F), 291.
Cladopbora Kiilz., 26, 55, 72, 102, 104, Cocconeina Ehrenb., 298, 299.
1(16, 291, 29S. cii'Spitosum (Kiitz.) nub., 299.
crispata (Roth.) Kilt.:., 105. Cistula Ehrenb., 2'.l'.l.
 Index 357
Cocconema cuspidatum (Kiitz.) nol>., Conochfete Klebahn, 52, 180, 181, 183.
299. comosa Klebuhn, 1S3 (fig. 71).
cymbifonne Ehrenb., 299. polytricha (Nordxt.) Kh'bnhn, 183.
Ehrenbergii (Kilt;.) nob., 299. Cooke, 1, 2, 91.
gracile (Jinbenh.) nob., 2'.i'.i.
Copepoda, 4.
hinceolatum Ehrenb., 299 (fig. Copeland, 126.
141 A). Corbiera Dang., 187.
prostratum (Berk) nob., 299. Corda, 223.
Cocconemacere, 291, 298 301. Correns, 244, 315.
Ccelastreas, 212, 213215. Coscinodiscacea?, 274, 276, 277.
Coelastrum NHg., 12, 25, 30, 212, 213. Coscinodiscus Ehrenb., 277.
cambricura Arch., 213 (fig. 87 A), lacustris Grim., 277 (fig. 127 D).
214. Cosmaridium Gay, 166.
cubicum Nii/j., 214. Cosmariese, 144, 14s, 150, 161.
distant Turn., 214. Cosniarium Corda, 137, 138, 144, 151,
microporum Nag-., 214. 166, 168, 169, 173.
proboscideum Bohliri, 214. abbreviatum Racib., 168.
})ulcltrnm Schmidle, 214. anceps Lund., 168.
reticulatum (Dang.) Seiui, 214 bioculatum Hre'b., 167 (fig. 62 I
(fig. 88). and J).
sphaericum Niitj., 213 (fig. 87 B Botrytis (Bon/) Men<'<jh., 168.
D), 214, 220. Cucurbita Brt'b., 168.
subpnlchnuii, Lagerh., 214. Dovrense Nordst., 168.
verrucosum Reinsch, 214. granatum Breb., 167 (fig. 62 B).
Coelosphserium N<i</., 311, 345, 346, 349. granatum var. subgranatum Nordxt. ,

Kiitzingianum \V'//.,349 (fig. 163 A).

J 167 (fig. 62 C and D), 168.
minutissimum Lemm., 349. Holmiense Lund., 16s.
Nii^eliaiuun Uiuji'i; 349. humile Gai/, 168.
Cohn, 3, 197, 307, 315. isthmiura We*t, 167 (fig. 62 M).
Colin'u'lht Schroder, 217. Klebsii Gutw., 139 (fig. 51 F).
Coleochaetaceff, 17, 30, 52, 66, 67 70, melauosporurn Arch., 168.
181. Meneghinii Breb., 167 (fig. 62 E
Coleochaate Breb., 12, 13, 54, 69, 181. G), 168.
irregularis. Prinijsh., 70. microsphinctum Nordxt., 168.
orbicnlaris Prinr/sh., 69, 205. moniliforme (Turp.) lialfx, 143.
pulvinata A. Br., 68 (fig. 17), 69. obliquum Nordst., 143.
scutata Breb., 67 (fig. 16), 69. ovale Ralfs, 168.
soluta Priiifjsh., 69. Pokornyauum (Gnat.) 1C. ((' G. S.
Coleospermum Kirchner, 322. West, 168.
Collection of Freshwater Algae, 7. prsemorsum Breb. , 167 (fig. 62 H),
Collemaceas, 325. 168.
Colletonema Breb., 292. pseudoconnatuni Nordst., 167 (fig.
Comere, 55. 62 X).
Conferva Lagerh. (and other authors), pygmamm Arch., 168.
81, 255, 256. pyramidatum Brt'b., 168.
aflinis Kiitz., 258. Ealfsii Breb., 168.
bombi/ciiia Ag., 257, 258. Eegnellii V'ilU; .143.
fontinalis Berk., 103. Eegnesii Reinsch, 167 (fig. 62 K
obsolete W. &
G. S. West, 258. and L).
jn'iiicilliformis Roth, 75. reniforme (Rulfs) Arc!/., 167 (fig.
Pllnil Dilleu., 256. 62 A).
rivuhtris Linn., 256. subcostatum Nordst., Ids.
ironnskioldii Flor. Dan., 75. subretusiforme W. <(<;. ,s. West, 168.
Confervales, 15, 28, 29, 30, 248, 249259. subspeciosuru Nordxt., 168.
Confen-iiid,;,-, 21, 27. subtuniidum Nonlxt., 168.
Conjufinta Yanch., 159. Cosmocladium Bi-efi., 138, 144, 151, 173.
Conjugate, 6, 15, 17, 21, 27, 30, 32, 33, coustrictuni Arch., 173 (fig. 66 A),
56,
50, 52, 54, 114178, 272. 174.
Conjugating-tube, 119. perissura Roy it-
Biss., 173 (fig.
Connecting-baud (of Diatom), 260. 66 C).
358 Index
Cosmocladium pulchellum Breb., 173 Cystocarp, 16, 35.
(fig. 66 B), 174. Cystococciis Nag., 202.
saxonicum De Bart/, 174. Cysts, 14, 102, 111, 121.
Cox, 265.
Craterospermum Braun, 121. Dactylococcopsis Hansr/., 346, 347.
Craticular state (of Diatoms), 270. montana W. < G. S. West, 348
Crucigenia Morr/'ii, 215, 216, 217. (fig. 162 A),
irregularis Wille, 217. rhaphidioicles Hansr/., 348.
quadrata Morren, 216 (fig. 90 D Dactylococcus Nag., 218, 223, 348.
and E). bicaudatus A. Br., 219 (fig. 91 A),
rectangularis (Nfiq.) Gay, 216 (fig. 223.
90 A C), 217. bicaudatus var. subramosus W. &
Tetrapedia (Kirchn.) W. cO G. S. G. S. H>Nf,219(tig. 91 B and C).
West, 216 (fig. 90 F). Debaryanus Beinsch, 219.
triangularis Chodat, 216. dispar W. G. S. West, 219 (fig.
<

Crucigenieee, 212, 215 217. 91 D).

Crustacea, 7, 145, 219, 270. infusionum Niig., 219, 220.
Cryoplankton, 5. Dactylothece Layer h., 30, 33, 246.
Cryptoglena Ehrenb., 32. Braunii Lagerh., 206, 246 (fig. 114).
Cryptonionadinacea?, 45. Dangeard, 51, 193, 214, 310.
Cryptomonas Ehrenb., 45. Dasygloea I'hwaites, 330, 332.
Cryptonemiaceas, 35, 43. amorpha Thwaites, 331 (fig. 151 C),
Cthonoblastus Kiitz. 332. ,
332.
Cultivation of Algte, 9. De Bary, 142.
Cyanophycece, 3, 306 352. Dabarya Wittr., 123, 124, 127.
Cyanophycin, 311. calospora (Palla) W. db G. S. West,
Cyanoplast, 308, 309. 53, 128 (fig. 46 B).
Cycas, 325. Desmidioides W. <f- G. S. West, 128
Cyclops, 219. (fig. 46 F I), 129, 143, 144.
Cyclotella Kiitz., 276. glyptosperma (De Bary) Wittr., 128
comta (Ehrenb.) Kiitz., 276. (fig. 46 A),
Kiitziugiana Chauvin, 276. lams (Kiitz.) W. & G. S. West,
Meneghiniana Kiitz., 276. 128 (fig. 46 C E).
operculata Kiitz., 276, 277 (fig. 127 Deiniga, 308.
B and C). Denticula Kiitz., 283, 285.
Cylindrocapsa Eeinsch, 54, 81, 82. elegans Kiitz., 283.
conferta West, 82 (fig. 26 E and tennis Kiitz., 283, 284 (fig. 130
F), 83. C and D).
geminella Wolle\&r. minor Hansq., Derbes & Solier, 257.
83. Desmagonium Ehrenb., 288.
involuta Heinscli, 82 (fig. 26 A D), Desmidiaceaa, 5, 6, 8, 11, 14, 16, 27, 28,
83. 30, 33, 50, 5254, 114116,
nuda Reinsnh, 83. 129, 135178, 266, 309.
Cylindrocapsaceae, 30, 66, 81 83. Desrnidiea3, 148.
Cylindrocystis Menegh., 143, 144, 148, Desmidium Ag., 143, 144, lol, 177.
149, 152, 155, 159. aptogonum Brei., 177 (fig. 69 D).
Brebissonii Menegh., 155 (fig. 54 cylindricum Grev., 140, 143, 177
H and 156.
I), (fig. 69 C), 178.
crassa De Bnry, 156. graciliceps (Nordst.) Lagerh., 178.
diplospora Lund., 142, 155 (fig. quadratum Nordst.. 177 (fig. 69 B),
54 J), 156. 178.
diplospora var. major West, 156. Swartzii Ag., Ill (fig. 69 A), 178.
Cylindrospermum Kiitz., 325, 326, 329. Desmonema Berk. < Thicaites, 322, 324.
catenatum Ralfs, 329. Wrangelii (Ag.) Born. ( Flah.,324.
iiKirritspeniittiit. Babenh., 329. De Toui, 72.
majus Kiitz., 321*. Diadesmis Kiitz., 292.
stagnate (Kiit:.) Jlont. <(' Flak., 328 Diatoma D. C., 284.
(fig. 150 E G), 329. elongatum A,/., 285 (fig. 131 A D).
Cylindrotheca Itabcnli., 278. hiemale (Lyngb.) Heib., 285' '(fig.
gracilis (Breb.) Grun., 279. 131 E F).
 Index
Diatoma hiemale var. mesodon (Kiitz.) Etiiloderniii <irncile De Toni, 205.
V. H., 285 (fig.
131 G). Endosphtera Klebx, 199.
vulgare Bon/, '285. Endosphseraceaj, 24, 30, 179, 197199.
Diatomacese, 260, 281, 284, 285. Engelmann, 315.
Diatomaceous Earths, 271. Enteromorpha Link, 30, 9 -I. 96, 97.
Diatomella Grev., 282. intestinalis (L.) Lint;, '.I? ifig. :'5 L).
Balfouriaua Grev., 283. Entocladia gracilis Ilmm/i., 20.1.
Diatomin, 2(J4. Entodesmis Borzi, 31.
Diatoms, 5, 260305. Epithemia Ureft., 298, 300.
Dichothrix /.miard., 337, 338, 339. Argus (Ehrenb.) Kiitz., 2(59, 300
interrupta W. G. S. West, 339
d> (fig. 142 D), 301.
(fig. 156 C), 340. Argus var. alpestris (If. ,S').)
Nordstedtii Born. &
Flah., 340. Rabenh., 301.
Orsiniana (Kiitz.) Born. iO Flah. gibba A'(V^., 300 (fig. 142 A), 301.
339 (tig. 156 D), 340. gibberula Kiitz. \a,r. producta Grim.,
Dictyocystis Lu<ierh., 235, 236. 301.
Hitchcockii (U'olle) Lanark., 236. tnrgida (Ehrenh.) Kilt:., 300 (fig.
Cleve, 293.
I>i<'ti/<>neis 142 B and C), 301.
Dictyosphffiriea?, 213, 235 238. Zebra (Ehrenb.) Kiitz., 301.
Dictyosphasrium Niig., 212, 235. Erernosphsera De Bary, 226, 227, 229.
Ehrenbergianum Niig., 236. viridis De Bary, 229 (fig. 99).
oviforme La<ierh., 236. Etard & Bouilhac, 311.
pulchellum Wood, 235 (fig. 104), Euachnanthes Scbiitt (sect, of Ach-
236. nanthes), 289.
reniforme BuJn., 236. Euastropsis Layerh.^ 20, 206, 209, 211.
'

Didyinopriiim Kiitz., 144, 177. Eichteri (Schmiille) L(i;/erli., 211

Dill, 22. 186. (fig. "86), 212.
Dillwyn, 1, 256. Euastrum Ehrenb., 137, 138, 144, 150,
Dimorphococcus A. Br., 33, 218, 221. 164, 211.
lunatus A. Br., 221 (tig. 93). ansatum Rulfx, 165.
Dinobryaceas, 31, 45, 47. binale (Turp.) Ehrenb., 143, 164
Dinobryon Ehrenb., 45, 47. (fig. 60 C), 165.
cylindricum Imhof, 48. crassum (Breb.) Kiitz., 165.
cylindricum var. divergens Lemm., Didelta (Turp.) Balfx, 142, 165.
47 (fig. 7 A and B). elegans (Breb.) Kiitz., 164 (fig. 60
elongatum Imhof, 48. A and B), 165.
protuberans Lemm., 48. gemmatum Breb., 165.
Sertularia'JSftrenft., 47 (fig. 7 C), 48. huraerosum Half*, 142.
sociale Ehrenb., 48. insigne Hass., 165.
Diplocolon AVVr/., 322, 324. oblongum (Grev.) Ralfx, 164 (fig.

Heppii AVny., 324. 60 D), 165.

Discoidese, 273, 274277. pectinatuni Breb.. 165.
Docidium Breb., 144, 150, 159, 162. verrncosum Ehrenb., 165.
iixpennn Breb., 153. Eudorina Ehrenb., 23, 30, 53. 194.
Baculum Brtb., 162 (fig. 57 A C). elegans Ehrenb., 194 (fig. 77), 195.
undulatum Bail., 162 (tig. 57 D Etulorhiellu Lemm., 194, 195.
and E). Eufrtii/iluria Ealfs (sectionof Fragi-
Dolichospermum Thwaites, 327. laria), 286.
Draparnaldia Bory, 87. Euglena, 23.
glomerata (rMf/i.) Ay., 87 (tig. 29), Ettmelosira Schtitt (sect, of Melosira),
88. 275.
plumosa (VauL-h.) Ay., 88. Eunai-icnla Schiitt (sect, of Navicula),
Drosera, 129. 293.
Dwarf-male, 61. Eunotia Ehrenb., 261, 288.
Dysphinctium Nag., 166 168. Air it* W. Sm., 288.
biceps (JP. Nm.) IK.I,., 289.
Edwards, 272. fle.rnoita var. liicapitiitii Grun., 289.

Ebreuberg, 265, 271. 'gracilis (Ehrenb.) Rubenh., 2ss (tig.

Elodea, 72, 205. 134 C), 289, 295 (tig. 137 F).
luuaris (Ehrenb.) Grun., 2s
1

Encyoiiema Kiitz., 299. , 1.

360 Index
Eunotia pectinalis (Kiitz.) Rabenh., 289. Glceocapsa Kutz., 13, 246, 345, 346,
^pectinalis var. undulata Ralfs,
288 350.

(fig. 134 A). Magma (Breb.) Kiitz., 350, 351

robusta Ralfs, 288 (fig. 134 B). (fig. 165 B).
tetraodon Elirenb., 289. polydermatica Kiitz., 350 (fig. 165
Veneris Kiitz., 289. C E).
Eunotiaceae, 281, 287289. punctata Nag., 350.
Ealfsiana (Hass.) Kiitz., 351.
Famintzin, 84. Glceochaste Lagerh., 344, 345.
Fischer, 308, 309. bicornis Kirchn., 345.
Fixcherella, (Born. & Flah.) Gom., 321. Wittrockiana Lagerh., 344 (fig.

Fishes, 270. 160), 345.

Flagellata, 23, 27, 33, 185, 248. Glceocystideas, 240, 244247.
Fleissig, 109. Glceocystis N>'in., 13, 19, 30, 242, 245,
Floridete, 3443. 246.
Fontinalis antipyretica, 4. innpla (Kiitz.) Kabenh., 246.
Foreliella Chodat, 4. gigas (Kiitz.) Lagerh.. 245 (fig. 113
Fossil Algte, 11. F H), 246.
Fragilaria Lijugb., 285.
infusionum (Schrank) W. <& G. S.
capucina Desmaz., 286 (fig. 132 C West, 245 (fig. 113 A E).
and D). vesiculosa Nag., 246.
construens (Ehrenb.) Grun., 286. Glosothece Ntig., 13, 346, 347.
Crotonensis (A. M. Echo.) Kitton, coufluens Nan., 347 (fig. 161 B).
286. liuearis Nag., 347 (fig. 161 A).
mutabilis (W. Sm.) Grun., 286. Glceotila Kiitz., 20, 26, 30,' 33, 75, 77, 78.
virescens Ralfs,r 286. protogenita Kiitz., 77 (fig. 22 C
Fragilariaceae, 281, 285287. E), 78.
Fragilarioidese, 263, 280-289. Glceotrichia J. Ay., 311, 337, 338, 340.
France, 22. natans (Hedw.) Rabenh., 89, 341.
Freeman, 198. Pisum (Ag.) Tlntr., 341.
Frenzel, 232. Glycogen, 311.
Fries, 75. Gobi, 251.
Fritsch, 84, 86, 312, 313. Golenkinia Chodat, 232, 233.
Frustule (of Diatom), 260. paucispinosa W. & G. S. West,
Fnattulia Kabenh., 294. 233 (fig. 102 F).
Fucoideee, 4449. radiata Chodat, 233 (fig. 102 D
Fungi, 10, 314. and E).
Gomout, 335.
GaiUionclla Bory (sect, of Melosira), Gomphoiieis Cleve, 297.
275. Gomphonella liabenh., 297.
Gametangia, 16. Gomphonema A<I., 103, 261, 297, 298.
Gametophyte, 18. acuminatum Elirenb. 298. ,

Gay, 20, 79, 89, 99, 103. constrictum Ehrenb., 297 (fig. 140
Geddes, 96. C), 298.
Geminella Turp., 26, 30, 75, 78. gemiuatnm (Lyngb.) Ag., 297 (fig.

interrupta Turp., 78 (fig. 23 A C). 140 A and B), 298.

Geuicularia I),- limy, 53, 144, 148, 149, parvulum Kiitz., 298.
153. Gomphonemaceae, 291, 297, 298.
elegans W. & G. S. West, 153. Gomphosphseria Kiitz., 346, 349.
Spirotsenia De Bary, 152 (fig. 53 I aponiua Kiitz., 349 (fig. 163 B).
and J), 153. Gonatonema Wittr., 54, 118, 122.
Gerassimoff, 123, 126, 132. Boodlei II'. t G. S. Wext, 118,
Gigartinacese, 35. 122 (fig. 45
F), 123. A
Girdle (of Diatom), 260. notabile (Has*.) Wittr., 123.
Girdle-view (of Diatom), 261. tropicurn W. tf- G. S. West, US.
Glaucocystaceaa, 30H. 309, 317. ventricosum Wittr., 118, 122 (fig.
Glaucocystidese, 3, 316, 317. 45 G
J), 123.
Glaucocystis Itziaxolin, 317. Gonatozygaa, 144, 149, 152.
Nostochinearum Itzigso.hn, 317. Gonatozygon De Bary, 137, 138, '144,
De Toni, 203. 148, 149, 153.
 Index 301

Gonatozygon Brebissonii De Bary, '

1.V2 Hazen, 86, 189, 256.
(fig. 53 A and B), 153. Hedgcock & Hunter, 40.
Brebissonii var. L-eve (Hilse) W. Hegler, 307311,
313.
<& G. S. West, 152 (fig. 53 C Helierella Bory, 159, 222.
-E). Helminthocladiea?, 36 40.
Brebissonii var. minutum W. ct Henfrey, 1.
G. S. Went, 152 (tig. 53 F Heribaud, 262.
and G), 153. Herposteiraceag, 30, 52, 66, 7072, 180.
Kinahani (Arch.) Rabenh., 153. Herposteiron Nfig., 52, 54, 70, 71, 72,
monotcenium De Bary, 152 (fig. 86, 89, 180.
53 H), 153. confervicola Nfig., 70 (fig. 18), 71
RalfsiiDe Bary, 153. (fig. 19 B D), 72.
Gongrosira Kiitz., 91, 111. ylobosa Nordst., 182.
stagnalis (West) Schniidle, 91, 92 pilosissirna (Schmidle) nob., 71 (fig.

(fig. 33 F).D 19 A), 72.

viriclis Kiitz., 91, 92 (fig. 33 A C). polychajte Hansq., 72.
Gonidangia 15. Heterocysts, 312, 313.
Gonidia, 15. Heterogamous gametes (or heteroga-
Gonimoblasts, 35. metes), 16.
Goniurn Miiller, 23, 30, 190, 191. Heterokonta?, 11, 29, 30, 33, 248259.
lacustre G. S. West, 191 (fig. 75 Hicks, 1.
B F). Hieronymus, 308, 310, 312, 317.
pectorale Miill., 191 (fig. 75 A), 192. Hildenbrandtia Nardo, 43.
sociale (Duj.) Warm., 192. rivularis (Liebm.) J. Ag., 43 (fig. 4).
Goroschaukin, 22. Himantidium auct. var., 288.
Grammatonema Kiitz., 285. Him, 57, 63.
Gray (S. F.), 135. Holacantluim (sub-gen, of Xanthidium),
Gregariniclte, 266. 169.
Grintzesco, 220, 230. Holocystis Hass., 165.
Grunow, 270. Holophytes, 13.
(Inininriii Rabenh., 302. Hormiditnn 18, 98.
Kiitz.,
Guano, 270. inrale Kiitz., 99.
ti

Gyyes Ehrenb., 159. piirietinum Kiitz., 99.

Gymnozyga Ehrenb., 144, 151, 178. Hormiscia Fries, 75, 76.
moniliformis Ehrenb., 177 (fig. 69 E Hormiscia Rabenh. Hansg. ; ;
De Toni,
and 178.F), 75, 76.
moniliformis var. gracilescens Hormococciis Chodat, 79.
Nordst., 177 (fig. 69 G). Hormogones, 15, 313.
Gyuogonidia, 191. Hormogoneas, 313, 318 342.
Gyrosigma Huss., 291, 292, 295. Hormospora Breb., 26, 30, 33, 73, 75,
attenuatum (Kiitz.) Rabenh., 296 77, 78, 81.
(fig. 138 A). mutabilis Breb., 77 (fig. 22 A).
Spencerii (Queck.) O.K., 296. ordinata W. &
G. S. West, 77 (fig.
22 B).
Htonatococcus Ag., 189, 206. plena Breb., 11.
imignis Hass., 206. Hormotila Borzi, 52, 184, 201, 205, 206.
Hallier, 265. mucigena Borzi, 205 (fig. 83 D).
Hansgirg, 1, 2, 18, 32, 126, 147, 205, 314, Huber, 72.
315, 327. Hyalotheca Ehrenb., 144, 151, 176.
Hantzschia Grim., 301, 302. dissiliens (Sni.) Bn'b., 143, 17G
Amphioxys (Ehrenb.) Grim., 303. (fig. 68 A D).
Hapalosipbon Nag., 38, 320, 321. mucosa (Dillw.) Ehrenb., 176.
Hibernicus W. & G. S. West, 321 neglecta Racib., 140, 176 (fig.
147), 322.
(fig.
68 E H).
intricatus West, 322. undulata Nordst., 176.
Haptera, 12, 52. Hyams & Richards, 307.
Hariotina Dang., 213, 214. Hydra viridis, 4, 230.
Hassall, 1, 206. Hydrianitm Rabenh., 200.
Hassallia Berk., 324. Hydrocoryne, 322.
Hauptfleisch, 12. Hydrocytium A. Br., 200.

235
362 Index

Hydrodictyaceae, 30, 180, 206209. Leibleinia Endl. (sect, of Lyngbya), 334.

Hydrodictyea?, 25, 207. Lemanea Bonj, 18, 20, 39, 40, 42.
Hydrodictyon Both, 11, 17, 25, 26, 30, catenata Kiitz., 41 (fig. 3 F).
206, 207. fluriatili*! Ag., 42.
reticulatum (L.) Lagerh., 208 (fig. parvula Sirod., 42.
Ml), 209. torulosa Kiitz., 41 (fig. 3 C and D), 42.
Hydruracese, 31, 45. Lemaneacese, 40.
Hydrurus Ag., 45, 46. Lemmerrnann, 1, 48, 194, 195, 217, 222,
fcetidus (Vill.) Kirchn., 46 (fig. 5). 224, 232, 252, 255, 272.
Hyphceotlirix Kiitz., 330, 335. Lemmermannia Chodat, 215, 216, 217.
Hypnocysts, 15. emarginata Chodat, 216.
Hypnospores, 15. Lemna, 72, 197, 198, 325.
gibba, 198.
Ichthyocercus W. & G. S. West, 144, minor, 198.
149, 150. trisulca, 198.
Tnactis Kiitz., 330. Leptosira Borzi, 92.
Ineffigiata W. oj G. S. West, 235, 237, Mediciana Borzi, 93.
238. Leptothrix Kiitz., 334, 335.
neglecta W. &
G. S. West, 238 (fig. Leucobryum glaucum, 247.
107). Leuronema Wallich, 175.
Infusoria, 267. Lewis (F. J.), 118.
Isoetes, 4. Lichen, 4, 314.
Isogamous gametes (or Isogametes), 16. Licmophora Ag., 283.
Isokontaj, 32. Limnactis Kiitz,., 340.
Itzigsohn, 314. Limnaja, 70.
Iwanoff, 86. peregra, 92.
Liparogyra Ehienb., 275.
Jackson & Ellms, 315. Lockwood, 269.
Joshua, 143. Lundell, 169, 171.
Jungermannia inflata, 4. Luther, 27, 29, 32.
Liitkemiiller, 136, 138, 148, 149.
Karsten, 32, 264, 269. Lychnis, 198.
Kieselguhr, 271. Lyngbya C. Ag., 307, 332, 333, 334, 335.
Kirchner, 32, 322. erugineo-co3rulea (Kiitz.) Gom., 334
Kirchneriella Schmidle, 25, 218, 225. (fig. 153 B
and C).
lunaris (Kirchn.) Mob., 226. jestuarii (Mert.) Liebm., 334.
obesa (West) Schmidle, 226 (fig. 96). major Menegh., 334 (fig. 153 A).
Kitton, 2(59. majuscula Harv., 316.
Klebahn, 51, 72, 181, 182, 269, 311. Martensiana Menegh., 334.
Klebs, 1, 9, 22, 23, 79, 111, 124, 136, ochracea (Kiitz.) Thur., 334.
138, 197, 199, 208, 259. Lyngbye, 256.
Klein, 196. Lyngbyeae, 330, 332 336.
Klercker, 20. Lysigonium Link (sect, of Melosira),
Kohl, 309, 310, 311. 275.
Kolkwitz, 132. Lysimachia, 199.
Kramer & Spiller, 271.
Kiihn, 54. Marquand, 1.

Kuntze, 222. Marx, 309.

Kutzing, 1, 75, 129, 206. Massart, 308, 309.
3Iastigonema Schwabe, 338.
Lagerheim, 1, 5, 20, 29, 31, 45, 51, 55, Mastigothrijr Kiitz., 338.
78, 80, 98, 99, 211, 236, 246, 256, Mastogloia Thwaites, 263, 292, 296.
257, 345. Dansei Thw., 297.
Lagerheimia Chodat. 25, 232, 234. Smithii Thw., 296 (fig. 139), 297.
gerievensis Chodat, 234 (fig. 103 Melanophycecs, 44.
A C). Melosira Ag., 261, 268, 272, 275.
subglobosa Lemm., 234 (fig. 103 D arenaria Moore, 275 (fig. 126 A
and E). and
B), 276.
Lateral conjugation, 119, 125. granulata (Elirenb.) Ralfs, 275,
Lauterborn, 262, 266. 276.
 Index 303
Melosira nummuloides (ttry) A;/., 275. Microcystis Kiitz., 346, 349, 350.
Koseana Rubenh., '275, 'JT'i. aeruginosa (Kiitz.) nob., 350.
varians A;i., 275 (tig. 120 C E). elabens (Breb.) Kiitz., 350.
MeloairacesB, 274 276. Flos-aquae (Wittr.) Kirclin., 350.
Mentha, 198. marginata M
<>.'////., 350 itig. 164 B).

Mereschkowsky, 264, 265. roseo-persicinus (Kiit:.) noli., 350.

Meridiou Ar/., 283, 284. stagnalis Lemm., 350 (fig. 164 A|.
circulare Ag., 284 (fig. 130 A andB). Microspora Tfeitr., 9, 2628, 100, 101,
circulare var. constrictum (Ralfs) 129.
V. H., 284. abbreviata (Rabenli.) Lmjerh., 101
Meridionacea?, 280, 281, 283, 284. (fig. 37 B and C).
Merismopedia Mei/en, 13, 310, 345, 346, amffina (Kiitz.) Lagerli., 101 (fig.
348. 37 A and F).
asruginea Breb., 348. amcena 101
var. crassior Haiisg.,
elegans A. Br., 348 (fig. 102 C). (fig. 37 E).
glauca (Ehrenb.) Nfig., 348 (fig. floccosa (ranch.) Thur., 101.
162 B). fontinalis (Berk.) De Toni, 103.
punctata Meyfn, 348. Lofgrenii Nordst., 28, 30.
Mesocarpese, 16, 18, 115, 117123. pachyderma (Wille) Lugerli., 101
Metacarpus Hass., 121. (fig. 37 D).
Mesotasniuin Nfig., 53, 138, 143, 144, Microsporaceae, 26, 27, 100 101.
148, 149, 154. Microsporales, 27, 30, 54, 5C>, 100 101.
caldariorum (Lagerh.) Hansg., 155. MicrothamniacesB, 30, 66, 89 93, 201.
chlamydosporum De Bary, 155 (fig. Microthamnion Xfig., 90, 201.
54 G). Kiitzingianum Nag., 90 (fig. 32
De Greyi Turn., 154, 155 (fig. 54 D). A D), 91.
Endlicherianum Niig., 155. strictissimum Rabenh., 90 (fig. 32
maerococcum (Kiitz.) Roy < Hi**., E), 91.
155 (tig. 54 E and F). ri'.i-ntar Cooke, 91.
maerococcum var. rnicrococcum Miquel, 268, 270, 271.
(Kiitz.) W. d- G. S. West, 154. Mischococcus Niii/., 28, 29, 250, 251.
purpureum W.d'G. S. West, 51,155. coufervicola Niiy., 252 (fig. 118).
violascens De liary, 51, 155. Mitzkewitsch, 51, 132.
Micrasterias Ag. (1827), 137, 138, 143, Mollusca, 270.
144, 146, 150, 165, 222. Monostroma Thur., 26, 30, 95, 96.
conferta Lund., 147. bullosa (Roth) Wittr., 96.
Crux-Melitensis (Ehrenb.) Hass., membranacea W. G. S. & West,
106 (fig. 61 A), 96, 97 (fig. 35 A Kl.
deuticulata Breb., 165, 16(i (fig. Monoticniece (sect, of Spirottenia), 154.
61 C). Moore, 229.
foliacea Bail., 146. Mougeotia Ag., 6, 72, 117119,
54,
furcata Ag., 147, 166. 121, 122124, 127.
Jenneri Rulfx, 166. calcarea Wittr., 121.
oscitans Ralfs var. mucronata capucina (Bory) Ag., 51, 120 (fig.
(Di.von) mile, 139 (fig. 51 C), 44 B).
166. elegantula Wittr., 121.
papillifera Breb., 165. gelatinosa Wittr., 122.
pinnatifida (Kiitz.) Ralfs, 166. genuflexa (Dillw.) Ay., 122.
rotata (Grev.) Ralfs, 165. gracillima (Haas.) Wittr., 120 (fig.
truncata (Corda) Breb., 165, 166 44 I), 122.
(fig. 61 B). parvula (Has*.) Wittr., 120 (fig.
Micrasterias Corda (1835), 222. 44 D H), 122.
fulcittti Corda, 222, 223. scalaris Has*., 114, 121.
Microchaate Thur., 322. viridis (Kiitz.) Wittr., 120 (fig. 44 C).
diplosiphon Go HI. var. Cumbrica Mougeotiopsis Palla, 127.
West, 323. Mounting Freshwater Algae, 8.
Microcoleus Desinuz., 330, 332. Movements of Diatoms, 264.
delicatulus W. <& G. S. West, 332, Miiller (O.), 262, 265, 266, 268, 300.
333 (fig. 152 A), Mueller, 256.
subtorulosus (Breb.) Gom., 332. Murray (G.), 20.
3(34 Index

Murray (J.), 42. Nitzschia hyalina Provasck, 264.

Myriopbyllum, 4, 340. leucosigma Benecke, 264.
Myxonema Fries, 19, 67, 84, 85, 86, 89. linearis (Ag.) W. Sm., 302.
amcenum (Kiitz.) Hazcn, 86. Palea (Kiitz.) W. Sm., 264, 302.
terme (Ag.) Rabenh., 86 (fig. 28). putrida Benecke, 264.
Myxophycese, 3, 4, 6, 8, 11, 12, 15, 17, sigmoidea (Ehrenb.) W. Sm., 300,
19, 32, 246, 306352. 302 (fig. 143 C and D).
Myxophycin, 308. sinuata (W. Sm.) Grun., 302.
Teema W. Sm., 279.
Nadson, 311, 317. Nitzschiaceas, 301 303.
Nageli, 19, 72, 209, 220, 265. Nitzschiella Eabenh., 302.
Nannandrium, 61. Nitzschiodese, 280, 301303.
Nardia emarginata, 4. Nodularia Mertens, 326, 328.
Navicula Bory, 262, 263 (transv. sect., sphaerocarpa Born. Flah., & 328
fig. 124), 264, 266, 267, 291, (fig. 150 H), 329.
292 293 spumigenia Mertens, 329.
alpina (ir! 8m.) Ralfg, 293 (fig. Nordhausen, 104.
136 A), 294. Nordstedt, 1, 28, 124, 143, 159, 182,
Amphisbama Bory, 269 (fig. 125 C). 222.
crasxiiit'rria Breb., 294. Nordstedt ia Borzi, 182.
cuspidata Kiitz., 263, 294. Nostoc Vauch., 4, 308 311, 313, 314,
S

exilis Gruii., 294. 325, 326, 327.

gallica (W. Sin.) V. H., 294. cceruleum Lyngbye, 326 (fig. 149 C),
lata Jlre'b., 294. 327.
limosa 269
Kiitz., (fig. 125 A). commune Vauch., 313, 327.
major Kiitz., 261 (fig. 123 B), 266, humifusum Carm., 327.
294. Linckia Borzi, 326 (fig. 149 A and
nobilis Ehrenb., 263, 266, 294. B).
nobilis var. Dactylus (Ehrenb.) V. macrosporum Menegh. , 327.
H., 261 (fig.' 123 A), microscopicurn Carm., 313.
perpusilla. (?;. 294. ,
muscoruin Ag., 327.
rhoinbdiilen Ehrenb., 294. piscinale Kiitz., 327.
serians (Brfb.) Kiitz., 293 (fig. punctiforme, 311.
136 D). sphasricum Vuuch., 327.
sphasrophora Kiitz., 293 (fig. 136 C). verrucosum Vauch., 327.
Kiitz., 263, 266, 269 (fig.
viridis Nostocaceje, 312, 313, 318, 319, 322,
125 D), 293 (fig. 136 B), 294, 324329.
295 (fig. 137 D and E). Notommata parasitica, 196.
Naviculaceas, 279, 291 297. Werneckii, 113.
Naviculoiderc, 279, 280, 291301. Nuphar, 4.
Nemalionaceffl, 35, 36. Nymphaoa, 4.
Nenia (Clausilia), 55.
Nephrocytium Niig. 25, 226, 228. ,
Occurrence of Freshwater Algaa, 3 7.

Agardhianurn Nag., 228. Odontidium Kiitz., 285.

ecdysiscepanum W. <& G. S. West, (Edogoniacese, 7, 14, 17, 27, 30, 52,
228 (fig. 98 B). 5765.
lunatum West, 228 (fig. 98 C E). (Edogoniales, 16, 30, 32, 33, 50, 55,
Nagelii Grun.. 228. 5765.
obe'sum W. & G. S. West, 228 (fig. (Edogonium Link, 12, 15, 17, 52, 54,
98 A). 55, 5759, 63, 65, 72, 256,
Netrium (Nag.) , 144, 148, 149, 156. 343.
Digitus (Ehrenb.) Itzigsh. <
Rothe, acrosporum De Bary, 65.
155 (fig. 54 K), 156. Ahlstrandii Wittr., 60 (tig. 12 D).
interruptum (Breb.) Liitkem., 156. Borisianum (Le Cl.) Wittr., 65.
oblongum (De Bary) Liitkem., 156. Boscii (Le Cl.) Wittr., 59 (tig.
Nitzschia //,-., 283, 301, 302. 11 A),
acicularis W. 302.
Sin., ciliatum (Hass.) Pringsh., 62 (tig.
coramunis Rabenh., 302. 14 C).
constricta (Kiitz.) Pritch., 302 (fig. cyathigerum Wittr., 62 (fig. 14 B).
143 A and B). giganteum Kiitz., 64.
 Index 305
(Edogoniuui Hirnii Gittic., 59 (fig. 11 B). Oscillatoria decolorata G. 8. West, 336.
Itzigsobnii DC Hnri/ var. minor irrigua Kiitz., 336 (fig. 154 B).
W,:<t, 60
(fig. 12 C), 05. lirnosa Ag., 336 (fig. 154 A),
lautumniarum Wittr., 61 (fig. 13 C princeps Vauch., 336.
and D), 65. splendida Grev. var. attenuata W.
obsoletum Wittr., 60 (fig. 12 A), &G. S. West, 336 (tig. 154 D).
platygynum Wittr., 65. tennis Ag., 336 (tig. 154 C).
punctato-striatum DC Banj, 65. Oscillatoriaceae, 199, 266, 312, 314
rufescens I17 (/-., 61 (fig. 13 A and (movements), 315, 318, 319,
B). 329336.
tapeinosporum Wittr., 04. Osterhaut, 35.
uudulatum (Breb.) A. Br., 62 (fig. Ott, 272.
14 A), 64. Ouracoccus Hass., 206.
zig-zag Cleve var. robustum W. &
G. S. West, 60 (fig. 12 B). Palla, 129, 309.
OligochtTtes, 145.
7, Palmella Lyngb., 19, 24, 30, 240, 243, 244.
Onderdonk, 265. hyalina Breb., 240.
Onychonema Wall., 143, 144, 151, 175. miuiata Leibl., 240.
filiformis (Ehrenb.)Eoy Biss., 175. & mucosa Kiitz., 240.
Nordstedtiaua Turn., 175 (fig. 67 Palmellaces, 5, 19, 22, 24, 25, 26, 30,
G and H). 180, 212, 239247.
Ooblastema filaments, 35. Palmellea:, 240 242.
Oocardium JVcV//., 144, 151, 174. Palmellococcus Chodat, 226, 229.
stratum Xiig., 173 (fig. 66 D F), 174. miniatus (Niig.) Chodat, 230 (fig.
Oocystideas, 213, 226 230. 100).
Oocystis Niig., 25, 226, 227, 228. Palmer & Keeley, 261.
asymmetrica W. <& G. S. West, 227. Palmodactylou Nag., 240.
crassa Wittr., 227 (fig. 97 C and D). subramosum N;ig., 241.
elliptica West, 227 (fig. 97 G). varium A7/V/.. 241 (fig. 108).
gigas Arch., 227. Palmodictyon Kiitz., 246.
panduriformis W. (- G. S. West, ( vifide Kiitz., 247 (fig. 115).
227 (fig. 97 E and F). Pandorina liory, 16, 23, 30, 192.
parva W. d G. S. West, 227. morum (Mull.) Bory, 193 (tig. 76
solitaria Wittr., 227 (fig. 97 A and A H), 194.
B).
Oodesmus Schmidle, 252. Paramecium, 4, 230.
Doedeiieinii Schmidle, 252. Partheuogonidia, 190.
Oogamous heterogamy, 16. Pectose, 51.
Oogonium, 16, 17. Pediastreaj, 25, 50, 197. 207, 209212.
Open nucleus (of Myxophyceas), 310. Pediastrum Meyen, 20, 25, 30, 206, 207,
Ophiocytiuir. Niig., 28, 253, 254, 256. 209, 212.
Arbuscula (A. Br.) Rabenh., 255 Boryanum (Turp.) Menegh., 210
(fig. 120 J). (fig. 85 F H; J L), 211, 220.
bicuspidatum (Borge) Lemm., 255. clathratiim Lemm., 211.
bicuspidatum forma longispina duplex Meyen, 210 (fig. 85 E), 211.
Lemm., 255 (fig. 120 H and I), glauduliferum Beiin., 210 (fig. 85 I).
capitatum Wolle, 255. integrum (Nil;/.), 210 (fig. 85 A), 211.
cochleare (Eichw.) A. Br., 255 (fig. pertiisum Kiitz., 211.
120 B G). simplex Meyen, 211.
graciliceps (A. Br.) Babenh., 255 tetras (Ehrenb.) Pnlfs, 209, 210 (fig.
(fig. 120 K). 85 C and D), 211.
majus Niig., 255 (fig. 120 A), tricornutum Borge, 210 (fig. 85 B).
parvuluru (Pert;/) A. Br., 255. Pedras uegras (of Angola), 306.
Ophrydium, 4, 230. Pellitan, 272.
Orthoneis Grun., 290. Peniejp, 144, 149,150, 157.
Orthosira Thwaites, 275. Penium, 137, 138, 140,
Br,'b., 144,
Oscillaria Bosc, 335. 148150, 157, 159, 160.
Oscillatoria ranch., 89,314 (movements), cucurbitinum Biss., 139 (fig. 51 D).
315, 329, 330, 332,333, 335, 336. curtum Breb., 158 (fig. 55 F).
acuminata Gotu., 336 (tig. 154 E). Cylindrus (Elirenb.) Breb., 157, 158
angustissima W. d G. S. West, 336. (tig. 55 A and B).
306 Index
Penium didymocarpum Lund., 141 (fig. Phymatodocis Nordst., 144, 149, 151.
52 D and E), 142. Phytheliea?, 213, 232234.
inconspicuum West, 157. Phythelios Frenzd, 232.
Libellula (Focke) Nordst., 144, 157, Pilinia Kiitz., 91.
158 (tig. 55 D). Piiiiinldrid.
Ehrenb., 262, 264, 266, 267,
margaritaceum (Ehrerib.) Breb., 157. 292.
minutissimum Nordst,, 157. Pithixcus Dang., 187.
minutum(JRai/s) Gleve, 144, 149, 158. Pithophora Wittr., 101, 107.
polymorphum Perty, 157. Keweusis Wittr., 107.
spirostriolatum Barker, 157, 158 (Edogonia (Mont.) Wittr., var. poly-
(fig. 55 C). spora Rendle & West f., 106
suboctangulare West, 158 (fig. 55 E). (fig. 41), 107.
subtile W. d- G. S, West, 138. Pithophoracese, 26, 30, 102, 106107.
Pennataa, 273, 279305. Placodermas, 144, 149, 156178.
Peridinieas, 4, 32. Plagiospermum Cleve, 121.
Peroniella Gobi, 29, 251. PlaiictonciiKi Schmidle, 78.
Pe talon ema Berk., 323. Plankton (Freshwater), 4.
Petit, 124, 272. Planogametes (or Zoogametes), 16, 17.
Pfitzer, 265, 272. Planorbis, 70.
Phacoteae, 186, 181190. Plectonema Thur., 330, 332, 333.
Phacotus Perti/, 190. Tomasiniana (Kiitz.) Born., 333.
lenticularis (Ehrenb.) Stein, 190. Pleodorina Xhaw, 195.
Phfeocapsaceae, 45, 48. Pleurenterium Lund, (subgen.), 171.
Phaaococcus Borzi, 31, 45, 48. Pleurocladia A. Br., 31, 45.
dementi (Menegli-.) Borzi, 48. Pleurococcaceff), 30, 83, 90, 179, 201206,
paludosus W. &
G. S. West, 48 212.
(fig. 8). Pleurococcus MenegJi.,1, 18, 26, 90, 201,
Phffiocystis Lagerh., 31, 45. 202, 203, 204, 206, 230.
Phasodactylon BoJilin, 31, 45. tus Nag., 230.
in in id

Phaeophycefw, 10,29,31,32,40, 44-49,264. nimbatus De Wild., 204.

Phaeophyll, 44. rut'escens (Kiitz.) Breb., 203.
Phfflosphsera W. & G. S. West, 31, 45, 49. rufescens var. sanguineus W. <(>
gelatinosa W. d- G. S. West, 49. G. S. West, 202 (fig. 81 B), 203.
Phseothamniaceffi, 45. vulgaris Menegh., 202 (fig. 81 A),
Phasothamnion Lagerh., 31, 45. 203, 204.
Phaeozoosporinaa, 44. Pleurosigma W. Sm., 295.
Phaeschizochlamys Lemm., 45. Pleurostauron Babenb., 294.
Philodendron, 55. Pleurotceniopsis (Lund.) Lagerh., 166.
Phormidium Kiitz., 330, 332, 333, 335. Pleurotasnium Nfig., 137, 138, 143, 144,
autumnale (.-I//.) Gom., 335. 150, 159,' 162.
molle (Kiitz.) Gom., 334 (fig. 153 D). coronatum (Breb.) Rabenh., 137,
purpurascens (Kiitz.) Gom., 335. 163 (fig. 58 A).
tenue (Menngh.) Gom., 334 (fig. Ehrenbergii (Breb.) De Bary, 163
153 E and F), 335. (fig. 58 B).
Phragmites, 340. maximum (Reinsch) Lund., 163.
PhycochromophycecB, 3, 306. nodosum (Bdil.) Lund., 147, 163.
Phycocyauin, 34, 308. Trabecula (Ehrenb.) Nfig., 163.
Pbycoerythrin, 34. truncatum (Breb.) Nfig., 163.
Phycophaein, 44. Podosira Ehrenb. (sect, of Melosira), 275.
Phycoporphyrin, 51. Polyblepharideae, 30, 186.
Phycoxanthin. 44. Polyblepharis Dung., 23.
Phyllobium Klebs, 198. Polychaetophora w'.d- G. S. West, 181, 183.
dimorphum Klebs, 199. lamellosa W. & G. S. West, 184 (fig.
Phyllosiphon, 13, 199. 72).
Alocasia3 Lagerh., 55. Polycystin; 308.
Arisari KiiJin, 54. Polycystis Kiitz., 349, 350.
maximum Ijiigerh., 55. Polyedrium Nag., 20, 231.
Philodendri Lagerh., 55. tetru'fdricum Nag., 231.
Phyllosiphonacefle, 109. Polymorphism, 18 21.
Phylogeny of Fresbwater Algae, 2133. Polytaniea (subgen. of Spirotaenia), 154.
 Index 307

Polytoma Ehrenb., 23. Rhaphidiuni Kiitz., 221, 222.

uvella Kln-i'iib., 23. aciculare A. Br., 223.
Porphyridium 340, 351.
X>'i<i., biplex Reinsch., 224.
cruenturn (Aff.) NH//., 351. conrolntum Rabenh., 224.
Porphyrosiphon Notarisii, 130. dupl,:r Kiitz., 223.
Potamogeton, 4. fiif<i-ii-nl<(titiii Kiitz., 223.
Prasiola Ag., 18, 30, 33, 98. var. xpirtili' (Turn.)
I'dxi-ii-iilitttim

crispa (Lightf.) Menegh., 99 (fig. Chod., 224.

36 D G"), 100. nirale Chodat, 223.
fiirfiii-iicen, Menegh., 100. Pfitzeri Schroder, 224.
parietina (Vauch.) Will,', 99 (fig. polymorphum Fresen. var. aciculare
36 A C). Rabenh., 223.
Prasiolaceas, 30, 98 100. polymorphumv&T. falcatinit Rabenh.,
Preservation of Freshwater Algae, 8. 223.
Pringsheim, 03, 69. polymorphum var. inirubili.-i W. &
Procarp, 16, 17, 34. G. S. West, 224.
Proliftra Vauch., 159. polymorphum var. npirale W. &
Protococcacese (or Autosporaceae), 1, 5, G. S. West, 224.
14, 25, 30, 33, '180, 201, 207, polymorphum var. tnmidum W. &
212238. G. S. West, 224.
Protococcoidea3, 11, 14, 15, 17, 1922, pyrngenum Chodat, 223.
30, 50, 52, 54. 56, 178247, 309. setigerum (Schrod.) W.
'
cfc G. S.
Protococcux Ag., 202,' 229, 230. West, 224.
Protoderma Kiitz., 12, 201, 204. xpirnle Turn., 224.
viride Kiitz., 205 (fig. 83 A C). Hhaphidonema Lagerh., 80.

Protomastigina, 23, 30, 31. nival e Lagerh., 80.

Pseudeunotia Grun., 2ss. Rhizoclonium Kiitz., 12, 26, 30, 72,
Pseudochffite W. 0. S. West, 88.
<( 102, 103, 129, 256, 343.
crassisetum W. & G. S. West, 89. _tl<i
riant* Rabenh., 103.
gracilis W. &
G. S. West, 88 hieroglyphicum Kiitz., 103, 104
30).
(fig. (tig. 39 A).
Pseudocilia, 51, 239, 243. hieroglyphicum var. Kochianum
104.
Pseudopleurococcus Snow, 202. (Kiitz.) Stock-in.,
var.
Pseudo-raphe (of Diatom), 263. hieroglyphicum riparium
Psilonemateas, 319 336. (Harv.) Stud; m., 104.
Pteromonas Seligo, 190. hieroglyphicum var. tortuosum
Pyrenoids, 12, 53. (Kiitz.) Stock-m., 104 (fig. 39
Pyxisporeae, 117.
B
E).
Kocliiiinnm Kiitz., 104.
Rabenhorst, 1, 3, 306. rivularis (L.) Kiitz., 256.
Eadais, 230. Rhizosolenia Ehrenb., 278.
Radiococcus Schmidle, 204, 212. eriensis Sin., 278.
nimbatus (Wild.) Schmidle, 204. longiseta Zach., 278 (fig. 128).
Radiotilurn Sclimidle, 26, 30, 33, 73, Bhizosoleniaceae, 278, 279.
75, 78, 81. Bhodophyceae, 10, 16, 17, 18, 32, 34
coujunctivum Schmidle, 79. 43, 93, 311, 351.
flavescens G. S. West, 78 (fig. Rhodymeuiaceas, 35.
23 D), 79. Rhoicosigma Grnn., 291.
Rafinesque, 159. Rhoicosphenia Grun., 263, 297, 298.
Balfsia O'Meara, 285. curvata (Kiitz.) Gntit., 298.
Raphe (of Diatom), 262. Richter, 55.
Richteriella Lemm., 232, 233.
Receptive spot, 17.
Red rain, 18'.). botryoides (Stock-in.) I.i'mm., 233
Red seaweeds, 34. '(fig. 102 A).
Red snow (plant), 5, 189. botryoides var. quadriseta (Li'inm.)
Beinxchii'lhi >.
setigera Schroder, 222 Chod., 233 (fig. 102 B and C),
224. 234.
Rendle & West f., 107. Rivularia (Both) Ag., 338, 340, 341.
Rhabd-odernut lineare Schmidle, 347. Biasolettiana Menegh., 340 (fig.
Rhabdonema arcuaturn (Ay.) Kiitz. 268. ,
157 A C).
368 Index
Eivularia dura Roth, 340. Schizochlamys delicatula West, 241 (fig.
haematites (D. C.) Ag., 340. 109 C).
minutula (Kiitz.) Born. & Flah., gelatinosa A. Br., 241 (fig. 109
340 (fig. 157 D and E). A and B).
Eivulariacese, 312, 313, 318, 337341. Schizogoniales, 26, 30, 33, 56, 98100.
Rosenvinge, 120. Schizogonium Kiitz., 98, 99.
18,
Rostafinski & Woronin, 259. crispuiii (Lightf.) Gay, 99.
Rotifer vulgaris, 100. murale Kiitz., 99.
Roy, 1. Schizomeris Leibleinii Kiitz., 76.
Roya W. d G. S. West, 138, 144, 150, Schizomycetes (or Bacteria), 3, 316.
158. Scliizonemn Ag., 292, 293.
Cambrica W. d G. S. West, 158 Scliiznjiliycefe, 3.
(rig. 55 I), 159. Schizophyta, 3.
obtusa (Breb.) W. & G. S. West, Schizosiphon Kiitz., 323, 338, 340.
159. Schizostauron Grun., 294.
obtusa var. montana W. & G. S. Schizothrix Kiitz., 330, 332.
West, 158 (fig. 55 G and H), calcicola (At/.) Gom., 331.
159. delicatissima W. d G. S. West, 33J.
Pseudoclosterium (Roi/) W. & G. S. fuualis W. d G. S. West, 331.
West, 158 (fig. 55 J and K), lardacea (Ces.) Gom., 331 (fig.
159. 151 B).
Eurnex, 198. Miillerii N<ig., 331 (fig. 151 A).
Schmidle, 1, 38, 40, 79, 92, 106, 183,
Sacheria Si rod., 40, 42. 204, 211, 216, 217, 237, 251,
fluviatilis (Ag.) Sirod., 42. 252, 346, 347.
fucina (Bory) Sirod., 41 (fig. 3E), 42. Schmula, 126.
rnamillosa Sirod., 41 (fig. 3 A and Schranim-id Dang., 344.
B), 42. Schroder, 12, 80, 138, 193, 222.
Saccodermae, 144, 149, 152156. Schroderia Lemm. 221, 222.,

Sachs, 3, 32, 306. setigera Lemm., 224.

Sauvageau, 313. Schulze (Max), 205.
Scalariform conjugation, 119, 124. Schulze's solution, 51.
Si-tilpruni Corda, 295. Schiitt, 32, 262, 272, 273.
Scenedesmus .!/<//<, 33, 218, 219. Sciddium A. Br., 254.
acutiformis Schrlkler, 221. Scirpus fluitans, 4.
acutus Meyeu, 220. Scotinosphffira Klebx, 199.
bijugatus (Turp.) Kiitz., 220 (fig. Scott, 111, 310.
92 C). Scytonema Ag., 4, 312, 314, 322, 323,
costatus Schniidle, 221. 324, 333.
denticulatus Lttgerh. var. linearis alatum (Berk.) Borzi, 323.
y^?(.sY/.,220'(fig. 92 I K), 221. Jigui'dtiim Ag., 323.
granulatus W. d
G. S. West, 221. mirabile (Dillw.) Thur., 323 (fig.
Hystrix Lagerh., 221. 148 A D).
obliquus (Turp.) Kiitz., 220 (fig. Myochrous Ag., 323.
92 A and B). Myochrous var. chorographicum
obtuisus Meyen, 220. W. d G. S. West, 306.
quadricauda (Turp.) Breb., 220 Scytonemacese, 312, 318, 319, 320, 322
(fig. 92 D-F). 324, 325.
quadricauda var. horridus Kirchn., Selenastrese, 213, 217226.
220 (fig. 92 G). Selenastrum Reinscli, 218, 225.
quadricauda var. maximus 17. & acuminatum Lagerh., 225 ffig. 95
G. S. West, 220 (fig. 92 H). E G).
spicatus W. d- G. S. West, 220 Bibraianum Reinsch, 225.
(fig. 92 L). gracile Jl<'i,,se1i, 225 (fig. 95 A D).
Scherffel, 63. Selenailenna Bohlin, 225.
Schiberszky, 265. Selenosphcerium Cohu, 215.
Schizacanthum Lund. (sect, of Xanthi- Senn, 48.
dium), 169, 170. Sexual organs, 16, 17.
Schizochlamys A. Br., 204, 240, 241, Sexual reproduction, 15 18.
242! Shaw, 195.
 Index
Siebold, 315. Spirogyra varians (Huxx. ) Kiitz., 125, 134.
Siphonea?, 21, 24, 2(5, 30, 33, 56,
11, velata Nordst., 127, 133 (tig. 49
101, 102, 108114, 199, 248. E G), 134.
Sirodot, 18, 20, 38, 40. Spirotaenia Hn'b., 55, 138, 144, 148,
Sirogonium Kiitz., 134, 135. 149, 154.
sticticnm Kiitz., 135. acuta Hihe, 138.
Siroxiphon Kiitz., 320. closteridia (l-in'h.) Arch., 154.
Smith (H. L.), 272. condensata Jin'b., 154, 155 (fig.
Solenoideffi, 274, 277279. 54 A).
Sorastrum Kiitz., 25, 30, 212, 215. obscura Haifa, 155 (fig. 54 ])).
Americanum (Bohlin) Schmidle, 215. truucata Arch., 155 (tig. 54 C).
spinulosum Nag., 215 (fig. 89). Spirotsenipse, 136, 144, 149, 154.
Spermatia (or Polliuoids), 16, 35. Spirulina Turp., 315, 330, 332, 333, 336.
Sphaerella Soiinnerf. 53, 187, 189. major Kiitz., 336 (fig. 155 B).
lacustris(Girod.) Wittr., 189 (fig. 74). siibfttilsd ffirsted, 336.
nivalis Somnicrf., 5, 189. tenuissima Kiitz., 336.
Sphaerocystis Chodat, 242. turfosa Jiuhi., 315, 336 (fig. 155 A).
Schroeteri Chodat, 242 (fig. 110), 243. Spondylosium lireb., 135, 142, 143, 144,
Sphcsrogonium Rostaf., 343. 151, 175.
Sphcerophora Hass., 275. nitens (U"//.) Arch., 140.
Sphasroplea A;I., 54, 108. papillatum II'. d: G. 8. West, 175
annuliua (liotli) A</., 108. (tin. 67 B), 176.

Sphseropleacese, 17, 26, 30, 50, 102, pulchellum Arch., 175 (fig. 67 C),
107108. 176.
Sphjerozosma Corda, 142, 143, 144, 151, Sporangia, 15.
174, 175, 251. Spores, 15.
excavatum Ralfn, 175 (fig. 67 D F). Sporophyte, 18.
granulatum lioy i Biss., 175. Squatnariaceffi, 43.
vertebratum Half*, 174, 175 (fig. Stapja Chodat, 243.
67 C). Staurastrum 3Ieyen, 137, 138, 144, 151,
Spheerozyga Ag., 327. 171.
Sphagnum, 198, 199, 325. acarides Nordst., 173.
contortum, 4. anatinum Cooke d) Wills, 172 (fig.
cuspidatum, 4. 65 A and B), 173.
Spirillum, 332. Arctiscou (Eh rnib.) Lund., 147, 173.
Spirochaate, 332. Aruellii Boldt, 173.
Spirocoleus Mobius, 334. brachiatum Ralfa, 172 (fig. 65 F).
Spirogyra Link, 12, 51, 53 55, 114, Brasiliense Nortist. var. Luudellii
115,117, 120, 123125, 127, W. d- G. S. IlVxf, 147.
134, 140, 256.
131, brevispinum lireb. 173.
calospora Clei'e, 134. capitulum Jin'b., 172.
communis (Hass.) Kiitz., 134. Cerastes Lund., 173
crassa Kiitz., 134. Dickiei Rail's, 141 (fig. 52 A C).
gracilis (Hans.) Kiitz., 134. elongatum Barker, 172 (fig. 65 E).
inflata (r<ntcli.) Rabcnh., 125, 133 furcigeruni Bn'b., 172 (fig. 65 G).
(tig- 49 D).
hexacerum (Khn'iib.) Wittr., 172.
majuscula Kiitz., 123, 131 (fig. 48 A), ineonspicuum Nurdxt., 148.
132, 134. iotanuui Wulle, 172.
maxima var. inrequalis Wollc, 126. jaculiferum Went, 147, 173.
/nirabilis (flaw.) Petit, 124. kjelmanui Will,-, 139 (tig. 51 E),
neglecta (Hass.) Kiitz., 132. 173.
nitida (Dillw.) Link, 132, 133 (fig. longispinum (I tail.) Arch., 147, 173.
49 A), 134. margaritaceum (Ehrerib.) Mi'iii'<i//.,
pellucida (Huss.) Kiitz., 132, 134. 172.
porticalis (1'uiich.) Cleve, 132. Ophiura Lund., 147, 173.
setiformis (Roth) Kiitz., 133 (fig. paradoxum Mii/i'ii var. longipes
49 B). Xord*t., 17:!.

Spreeiana Rnbenh., 133 (fig. 49 C). pelagicum M'. a- (i. S. West, 173.
tenuissima (flx.s.) Kiitz., 125, 131 pileolatum I>r<'b., 172.
(tig. 48 C), 134. polytrichum Perti/, 172 (tig. 65 D).
370 Index
Staurastrura pseudopelagicum II'. d G. Surireila robusta var. splendida (Ehrenb.)
S. West, 173. V. H., 304, 305 (fig. 145 C).
punctulatum Brel., 172 (fig. 65 C). spiralis Kiitz., 304.
pygmaeum Brfb., 172. Surirellacese, 280, 303305.
teliferum Ralfs, 172. SurirelloideaB, 280, 303305.
tumidum Breb., 172. Symphyosiphon Kiitz., 323, 338.
verticillatum Arch., 173. Symploca Kiitz., 332, 334.
Staidw/enia Kiitz., 215, 216. muralis Kiitz., 333 (fig. 152 L and
Stauroneis Ekrenb., 292, 293, 294. C), 334.
acnta W. Sm., 293 (fig. 136 F). Syncrypta Ehrenb., 47.
Phcenicenteron (Nitzsch) Ehrenb., Volvox Ehrenb., 47.
293 (fig. 136 E), 294. Synechococcus Nfig., 343, 345, 346, 347.
Stauros (of Diatom), 262. aaruginosus Nan., 347. .

Staurosira Ehrenb. (sect, of Fragilaria), major Schroet. 347 (fig. 161 D and E).
,

286. Synedra Ehrenb., 265, 272, 286.

121.
StduroxfU'DitiDii- Kiitz., Acus (Kiitz.) Grun., 287.
Stephanodiscus Ehrenb., 276. biceps W. Sm., 289.
Hantzschianus Grun., 277 (fig. capitata Ehrenb., 287.
127 Aj. pulcbella Kiitz., 286 (fig. 132 A and
Stephanokontce, 32. B).
Stephanosphaera Colin, 192. Ulna (Nitzsch) Ehrenb., 28(5.
pluvialis Colin, 192, 193 (fig. 76 K). Ulna var. splendens (Kiitz.) V. H.,
Stichococcus Nag., 13, 19, 26, 33, 55, 287.
75, 79. Syngeneticre, 15, 17, 44 49.
bacillaris Nag. 79 (fig. 24 A), 80. Synura Ehrenb., 45, 46.
dissectus Gay, 79 (fig. 24 C), 80. Uvella Ehrenb., 46 (fig. 6).
flaccidus (Kiitz.) Gay, 79 (fig. 24 B),
80. Tabellaria Ehrenb., 261, 281.
variabilis W. d- G. S. West, 79 (fig. fenestrata (Lyngb.) Kiitz., 282 (fig.
24 D), 80. 129 D and
E).
Stichoglcea Chodat, 31, 45, 49. fenestrata var. asterionelloides
olivacea Chodat, 49. Grun., 282.
Stigeoclonium Kiitz., 84 86. flocculosa (Eoth) Kiitz., 200, 282
Stigonema An., ' 4, 311, 312, 314, 31,7, (fig. 129
F and G).
320. Tabellariaceae, 280, 281283.
oompactum var. Brasiliense Wille, Tardigrades, 7, 145.
811. Temnogametacea3, 116, 135.
hormoides (Kiitz.) Born & Flah., Tetmemorus Ralfs, 138, 144, 150, 159,
321. 163.
informe Kiitz., 321. Brebissonii (Meneyh.) Ralfs., 164.
mamillosum Ag., 321. granulatus (Breb.) Ralfs, 163 (fig.
minutum Haas., 320 (fig. 146 A 59), 164.
and B). lasvis 164.
(Kiitz.) Ralfs,
ocellatum (Dilhv.) Tliur., 320 (fig. Tt'trafliastrum Dixon, 165.
146 C E). Tetracoccus West, 204, 235, 236, 237.
Stigonemaceae, 312, 313, 318, 319322, botryoides West, 236 (fig. 105), 237.
325. ninibatus Scbmidle, 204.
Stipitococcus II'. G. S. Went, 29, 250.
d'-
Tetracyclus Ralfs, 281.
urceolatus W. < G. S. West, 250 lacustris fialfs, 281 (fig. 129 A C).
(fig. 116), 251. rupestris (^1. Br.) Grun., 281.
Stizenberger, 3, 306. Tetraedrieae, 213, 231232.
Stockrneyer, 103, 308, 309. Tetraudron Kiitz., 20, 231, 34'..
Stomatochytrium Cunn., 198. caudatnm (Corda) Haitsg., 231 (fig.
Streptonema Wall., 144, 149, 151. 101 B).
Suriraua, 304. enorme (Ralfs)'
Hans a. ,
231 (fig.
Surireila Turp., 303, 304. 101 D).
biseriata Breb., 303, 305 (fig. 145 A), horridum II'. < G. S. West, 231
linearis If. Sm., 305 (fig. 145 B). (fig. 101 EG).
ovalis Breb., 304. minimum (A. Br.) Hansn., 231 (fig.
robusta Ehrenb., 304. 101 A).
 Index 371

Tetraedron pentae'drica 1C. <(' G. S. Triclionimx Allman, 327.

Wext. 216. Trinema acinus, 100.
regulare Kiitz., 231 (fig. 101 C). Triploceras Bail., 144, 149, 150.
Tetragon! urn W. ct G. S. West, 191. Tripoli, 271.
Zacus/re W. A G. S. West, 192. 'I'n.chiscia Kiitz., 90, 201, 202, 203.

Tetrapedia AV/H.-V/I., 216, 343, 345, 3411, aciculifera (Lmjerli.) I In UK;/., 204.
348. aspera (Reiuxcli) llmi*;!., 203 (fig.
glaucescens (ir///>-.) 349.
/.W./f. 82 A F), 204.
morsa II'. ,< G. S. Wext, 216. birta (Reinxch.) Hansy., 203 (fig.
Reinscbiana 4rcfc., 348 (fig. 102 D), s2 G H), 204.
349. paucispinosa West, 203 (fig. 82 I
setigera Arch., 349. and J).

Tetraspora Link; 13, 24, 2G, 29, 30, 51, reticularis (lli'hisrli) Ilunxi/., 203
53, 55, 239, 240, 243. (fig. 82 K i,204.
explanata A//., 243. Tn/blion.'lla W. Sin., 302.
gelatiuosa ( l'<inch.) l>cxr., 243. Tunicates, 270.
lacustrix Lemm., 242, 243. Turbellarians, 7.
lubrica (Roth) A;/., 243 (tig. 111). Turner, 91, 140.
Tetrasporese, 240, 243, 244. Turpin, 78.
Tetraspores (or tetragonidia), 4.
Tetrastrum Chodnt, 217. Ulothrix Kiitz., 9, 12, 20, 30, 73, 74, 75,
beteracantbum (Xordst.) Chad., 217. 77, 79, 80.
staurogeniaeformis (Sclirml.) Chad., aequalis Kiitz., 76 (tig. 21 F). A
2ir, (tig. 90 G and H), 217. sequalis var. catseniformis (Kilt:.)
Thamniochsete Gy, 89. llubcnh., 76 (tig. 21 G).
acnleata W. G. S. West, 89
(
:
-

(fig. monilifonnis Kiitz., 76 (tig. 21 H

31). and I).

Hu.beri Gay, 89. rmlii-nitx Kiitz., 99.

Thorea .Bon/, 40. subtilis Kiitz., 73, 74 (fig. 20 C F),
ramosissima Bory, 40. 76.
Thuret, 1, 239. subtilis var. variabilis (Kiit:.)
Tilden, 85. KirrJm., 76.
Timberlake, 2(is. zonata (Web. et Mohr) Kiitz., 73,
Tolypothrix A'/rt,:., 308, 311, 312, 318, 74 (fig. 20 A and B), 75, 76.
322, 324. Ulotrichaces;, 13, 15, 16, 17, 19, 25, 26,
lanata (Desv.) Wartin., 323 (fig. 30, 33, 66, 7381, 83, 252.
148 E), 324. Ulotrichales, 66.
tenuis Kiitz., 324. Ulva, 26, 95, 96.
pygmsea Kiitz., LS4. Ulvacea?, 16, 26, 30, 9597, 180.
Treutepoblia Mart., 4, 93, 95. Ulvales, 26, 30, 56, 9597, 98.
aurea :,I,trt., 94 (tig. 34 A C), 95. Urococcus Kiitz., 206.
calamicola (Zell.) De Toni, 94 (tig. iusignis (Hass.) Kiitz., 206.
34 D F), 95. Uroglena Ehrenb., 45, 47.
odorata (A;i.) Wittr., 95. Volvox Ehrenb., 47.
itmbriiin (Kiitz.) Born., 95. Urouema Lagi'i-li., 75, 80.
Trentepohliaceaj, 16, 30, 66, 67, 90, confervicolum Lagerh., 80.
93-95, 181. Urosporu Arescb., 75.
Tribonema Verb. & Sol., 9, 28, 81, 100, I'rxincUo Turp., 159.
253, 255, 256. Utricularia, 4, 93.
affine (Kiit:.) nob., 258. minor, 322.
bombycinum (Afi.) Derb. <(' Sol.,
257 (tig. 121 A G), 258. Vacuolaria, C'ii'iih.. 29, 30.

bombycinum forma minor (II7//t') Yaginarieffi, 330 332.

nob. 257 (tig. 121 H and I), 258.
, Valoniaceae, 102, 108.
obsoletum nob. 258. , Valves (of Diatom), 260.
Tribonemaces>, 28, 30,249,252, 253258. Valve-view (of Diatom), 261.
Trichogyue, 16, 17, 34. Van Heurck, 272.
Tiichome (of Myxophycese), 31 8. Vanbeurckia lirvb., 2li3, 291, 292, 294.
Trichopbilus Weber. 55. rbomboides (Ehrenb.) Hn'.b., 294,
Trichopboreffi, 319, 337 342. 295 (tig. 137 A and B).
372 Index
Vanheurckia rhomboides var. saxonica Wildeman (de), 94, 204.
G. S. West, 294.
(liabi'iili.) Wille, 1, 22, 28, 32, 59, 72, 73, 99,
vulgaris (Thw.) V. H., 294. 103, 170, 171, 189, 202, 217,
Vaucheria D. C., 14, 15, 17, 55, 109, 310, 311.
111, 113, 248, 291, 339, 343. Willed Schmidle, 215, 217.
a versa Haas, 114. Wisselingh, 51, 132.
dichotonia (Lyngb.) Ay-., 114. Wittrock, 1, 63, 80, 81, 107, 118, 119.
geniinata (Vauch.) D. C., 110 (fig. Wolle, 1. 2, 18, 77, 182, 314.
42 A, F H), 113. Wright, 316.
hamata (Vauch.) Lyngb., 112 (fig.
43 C and D), il4. Xanthidium Ehrenb., 138, 144, 151, 168,
sericea Lynijb., 110 (fig. 42 B and 170.
D), 112 (fig. 43 E), 114. antilopaeum (Breb.) Kiitz., 169 (fig.
sessilis (Vauch.) D. C. 55, 110 (fig. 63 B), 170.
42 C and
E), 112 (fig.
43 A armatum \Iirfli.) liabciih., 169 (fig.
andB), 113. 63 A), 170.
synandra, 112. conciunuin Arch., 170.
terrestris Lyngb., 114. cristatum Breb., 170.
Vaucheriaceae, 1517, 27, 29, 30, 33, Xanthophyll, 248, 264.
50, 53, 109114. Xenopus lavis, 7.
Vaucheriales, 29, 248.
Vegetative multiplication, 13, 14. Zacharias, 308, 309, 310.
Voigt, 270. Zonal-view (of Diatom), 261.
Volvocacec-e, 5, 16, 23, 24, 30, 51, 110, Zoiiatriciiiit J. Ag., 340.
179, 184197, 202. calcarea (Eng. Bot.) Endl., 340.
Volvocea?, 186, 190197. Zoogonidia, 15.
Volvox (L.) Ehrenb., 17, 23, 30, 190, Zoospores, 15.
195, 196, 197. Zopf, 308.
aureus Ehrenb., 196 (fig. 78 A, C, Zukal, 308, 309.
and D), 197. Zumstein, 23.
globator (L). Ehrenb., 196 (fig. Zygnema Ag., 51, 54, 115, 117, 123
7H B), 197. 125, 127, 129, 130, 140.
anomalum (Hass.) Conke, 131.
Wager, 258, 308, 310. cruciatum (Vauch.) Ag., 131.
Wallich, 140. ericetorum (Kiitz.) Hansg ., 129, 130,
Water-bloom, 315, 316. (fig. 47 C).
Water-net, 209. insigne (Haxx.) Kiitz., 130 (fig.
Weed, 307. 47 E), 131.
Wellheim (Pfeiffer E. v.), 8. leiospermum DC Ban/, 130 (fig.
Welwitsch, 130. 47 D).
West (G. S.), 2, 6, 27, 28, 55, 72, 138, pachydermum West, 28, 30.
143, 145, 146, 148, 168, 205, pachydermum var. coufervoides
307, 314. Went, 117.
West (W.), 117, 125, 126, 140, 170, pectinatum (!'?/<//.) A</., 131.
221, 236, 254. Ealfsii (Hass.) !>< Jinn/, 130 (tig
West (W.) fc West
(G. S.), 5, 2628, 47 F).
53, 115, 117, 118, 123, 125 spontaneum Nordxt. , 30, 123.
127, 129, 131, 138, 140, 141, stellinum 130 (fig
(Vauch.) A;/.,
143, 146, 183, 192, 214, 216, 47 A).
221, 233, 237, 250, 253, 282, Vaucherii Ag. var. stagnate (Hass.)
3U6, 312, 321, 324. Kirchit., 130 (fig. 47 B), 131.
}\','*trU<, J)c Wild.,
204, 236. Zygnemaoeae, 7, 11, 13, 14, It;, 30, 50,
iiimbdttix ]>e Wild., 204. 54, 114, 115, 116135.
Whipple, 186. Zygnemeaj, 115117, 119, 121, 123
Whipple & Jackson, 270. 135, 143.
W hippie & Parker, 13. Xi/<i<>t/(iinin Kiitz., 124, 129.

CAMBRIDGE: PRINTED BY j. AND c. F. CLAY, AT THE UNIVERSITY PRRSS.