FROM THE LIBRARY OF
WILLIAM A. SETCHELL,i864-i943
PROFESSOR OF BOTANY
BULLETIN
ILLINOIS STATE LABORATORY
NATURAL HISTORY
URBANA, ILLINOIS, U. S. A.
VOL. VIII. MAY, 1908 ARTICLE I.
THE PLANKTON OF THE ILLINOIS RIVER, 1894-1899, WITH
INTRODUCTORY NOTES UPON THE HYDROGRAPHY OF THE ILLINOIS
RIVER AND ITS BASIN. PART II. CONSTITUENT ORGANISMS AND
THEIR SEASONAL DISTRIBUTION.
BY
C. A. KOFOID, PH.D.
< -2-
B1OLOSY
LIBRAIY
BULLETIN
ILLINOIS STATE LABORATORY
NATURAL HISTORY
URBANA, ILLINOIS, U. S. A.
VOL. VIII. ARTICLE I.
THE PLANKTON OF f THE ILLINOIS RIVER, 1894-1899, WITH
INTRODUCTORY NOTES UPON THE HYDROGRAPHY OF THE ILLINOIS
RIVER AXD ITS BASIN. PART II. CONSTITUENT ORGANISMS AND
THEIR SEASONAL DISTRIBUTION.
BY
C. A. KOFOID, PH.D.
THE ILLINOIS PRINTING COMPANY
DANVILLE, ILLINOIS
BIOLOGY LIBRARY
CONTENTS.
PAGE.
Introduction i_i 7
Distribution of Collections by Months (Table) 4
Methods 5
Acknowledgments 7
Definitions 7
The Composition of the Plankton 10
Comparison of Fresh-water and Marine Plankton 12
Organisms of the Plankton 13
Constituent Groups of the Annual Plankton of the Illinois River (Table) 16
Discussion of the Statistical Data of the Species composing the Plankton
of the Illinois River in 1894-1899 18-290
Cryptogamia ; 18-62
Bacteriaceae 18
Schizophyceae 19
Discussion of Species 20
Chlorophyceae 22
Discussion of Species 24
Bacillariaceae 34
Factors controlling Diatom Production 35-43
Diagram showing the seasonal distribution of diatoms, total
plankton, nitrates, and thermograph and hydrograph of Illi-
nois River at Havana for 1898 37
Discussion of Species 43
Conjugatae 59
Discussion of Species 60
Phanerogamia . 62
Protozoa 62
Mastigophora 63
Location and Amplitude of Pulses of Chlorophyll-bearing Organ-
isms in the Illinois River (Table) 64
Discussion of Species 68
Rhizopoda 92
Discussion of Species 95
Heliozoa 113
Discussion of Species 113
Sporozoa 114
Ciliata 115
Discussion of Species 118
Suctoria 131
Discussion of Species 131
Porifera.. 132
SZ246526
PAGE
Coelenterata 132
Platyhelminthes 133
Turbellaria 133
Trematoda 134
Cestoda 135
Nemertini 135
Nematelminthes 135
Nematoda 135
Acanthocephala 136
Annulata 136
Oligochaeta 136
Naididag 137
^Eolosomatidag 137
Rotifera 138
Rhizota 139
Discussion of Species 140
Bdelloida 141
Discussion of Species 141
Ploima 144
Pulses of Ploima (Table) 146
Discussion of Species 147
Tabular statistical data :
Pulses of Anurcea cochlearis ISO, 153
Pulses of A nurasa hypelasma 155
Pulses of Asplanchna brightwellii 156
Evidence of polycyclic character of seasonal distribution of above . 158
Evidence of same for Asplanchna priodonta 161
Pulses of Brachionus angularis 165
Collection data for Brachionus bakeri and varieties 169, 170
Pulses of Brachionus budapestinensis 176
Pulses of Brachionus militaris 179
Pulses of Brachionus mollis 181
Brachionus pala and varieties. Average number per collection . 182
Pulses of Brachionus pala and varieties 184
Brachionus pala, type form. Sexual cycle 188
Seasonal distribution of B. pala and its var. amphiceros 189
Seasonal limits of B. pala var. dorcas 191
Pulses of Brachionus urceolaris 193
Brachionus urceolaris and varieties and B. variabilis. Average
number per collection 196
Pulses of Polyarthra platyptera 204
Pulses of Synchceta pectinata 211
Pulses of Synch&ta stylata 213
Pulses of Triarthra longiseta 218
Scirtopoda 219
Seasonal fluctuations of Pedalion mirum (Table) 220
Gastrotricha . . 221
PAGE
Entomostraca 221
Branchiopoda 222
Cladocera 223
Cladocera and Hydrographic Fluctuations (Table) 223
Discussion of Species 225
Tabular statistical data:
Bosmina and hydrographic fluctuations . r- 228
Seasonal distribution of Chydorus. Average number per m. 3 . . . 235
Effect of temperature on numbers of Chydorus 238
A typical pulse of Daphnia cucullata 242
Diaphanosoma population, with data of temperature and river
level 248
Moina and hydrographic changes 254
Ostracoda 257
Discussion of Species 258'
Copepoda 258
Copepoda and Hydrographic Changes (Table) 260
Discussion of Species 261
Cyclops viridis var. insectus and Hydrographic Changes (Table). . 274, 275
Relative numbers of adult, young, and larval Cyclopidae (Table). . 278
Amphipoda 283
Arachnida 283
Acarina 283
Tardigrada 284
Hexapoda 284
Ephemerida 285
Hemiptera 285
Diptera 285
Mollusca 287
Gastropoda ' 287
Lamellibranchiata 287
Bryozoa 288
Discussion of Species 288
The Periodicity in the Multiplication of .the Organisms of the Plankton. . . 291-311
Comparison of Plankton Pulses and Lunar Cycle (Table) 296-299
Gaseous Contents of Pond Water (Table) 306
Relation of Pulses of Chlorophyll-bearing Organisms to Lunar Cycle
(Table) 308
General Considerations on Seasonal Changes 312
Lake versus River Plankton 312
Organisms per Cubic Meter in Plankton of Illinois River in 1898 (Table) . . 314-340
Bibliography 341-354
Explanation of Plates 355
Diagrams of Seasonal Distribution 356360
Errata and Addenda. . 361
ARTICLE I. Plankton Studies. V. 1 The Plankton of the Illi-
nois River, 1894-1899. Part II. Constituent Organisms and their
Seasonal Distribution. BY C. A. KOFOID.
INTRODUCTION.
This paper gives the results of a statistical study of a series of
quantitative plankton collections made in the channel of the Illinois
River near Havana, 111., at the Illinois Biological Station, in 1894-
1899. The environmental conditions and the volumetric results of
this investigation have been given in Part I. (Kofoid, '03), published
in Volume VI. of this Bulletin.
Of the 235 collections made in channel waters and used in the
quantitative study, only 182 were subjected to numerical and
qualitative analysis. The omitted collections were intercalated at
brief intervals of one to several days between those enumerated,
principally in the summer of 1895 and during the winter flood of
1896 and the summer of the same year. The collections chosen for
this study, whenever possible, represent a weekly interval, and a
full list of all collections, with environmental data, may be found in
Table III. of Part I. The chronological distribution of the collec-
tions studied by the statistical method is given in the table on
the following page.
The work of enumeration and the primary tabulation was com-
pleted at Urbana December 31, 1900, when my formal connection
with the State Laboratory ceased. The manuscript has been
1 The four preceding numbers of this series, all by the present writer, have
been published as articles of the Bulletin of the Illinois State Laboratory of
Natural History, as follows:
Article I., Vol. V. Plankton Studies. I. Methods and Apparatus in Use in
Plankton Investigations at the Biological Experiment Station of the University of
Illinois.
Article V., Vol. V. Plankton Studies. II. On Pleodomna illinoisensis , a New
Species from the Plankton of the Illinois River.
Article IX., Vol. V. Plankton Studies. III. On Platydorina, a New Genus
of the Family Volvocidce, from the Plankton of the Illinois River.
Article II., Vol. VI. Plankton Studies. IV. The Plankton of the Illinois
River, 1894-1899, with Introductory Notes upon the Hydrography of the Illinois
River and its Basin. Part I. Quantitative Investigations and General Results.
prepared at Berkeley, being completed in May, 1904, after my
connection with the University of California was begun. My
separation from the collections and the library of the State Labora-
tory has rendered impossible some verifications, comparisons of
specimens with more recent literature, especially among the algae,
DISTRIBUTION OP COLLECTIONS BY MONTHS.
'94.
91
'96.
'97.
'98.
'99.
I
4
3
5
II
1
4
2
4
4
Ill
5
1
5
4
IV
2
4
1
4
V
4
1
5
VI
2
1
5
1
4
VII
2
4
5
3
4
VIII
1
5
6
4
5
IX..
2
4
2
4
4
X
1
5
1
4
4
XI
1
4
1
5
5
XII
1
5
2
4
5
Total
10
31
43
30
52
13
some desirable amplifications from omitted intermediate collec-
tions, and the elimination of a few minor errors in the statistics.
It should be understood that the data of this paper are derived
from channel collections, and the conclusions apply only to that
region. Conditions of plankton development in the adjacent back-
waters, as shown in Part I., differ greatly in volumetric character
and seasonal distribution. The composition of the plankton and
the seasonal distribution of its constituent organisms also exhibit
there many points of difference from those here described for channel
waters.
METHODS.
The collections were preserved in bottles of uniform capacity
(60 cm. 3 ), in alcohol-formalin mixture (2 per cent, formalin in 70
per cent, alcohol), and after measurement by the centrifuge were
released from the compressed condition in the measuring tubes and
returned to the containers.
The counting was done by a modified Sedgwick-Rafter method
(see Kofoid, '97), in which 1 cm. 3 of a suitably diluted plankton is
distributed evenly in a cell 20 X 50 mm. The plankton was diluted
or condensed (from 60 cm. 3 of fluid) according to the quantity of
plankton and the amount and nature of the silt. Larger organisms
such as the Entomostraca were counted in the whole catch, or in
larger collections in Yio to 1 / 50 of the total catch ; and the smaller
organisms in l l^ to V^o- The filter-paper catches which supple-
mented those of the plankton net from August 3, 1896, to the end
of the series, March 28, 1899, were often subjected to considerable
dilution on account of the great amount of fine silt in the collections,
from Vio "to Yioo being the limits of dilution as a rule.
The even distribution of the organisms in the Rafter cell was
secured by shaking the collection in a mixing cylinder gently till
the sediment was thoroughly distributed, and taking the sample
immediately with a long 1 cm. 3 pipette, inserted to the bottom of
the jar and raised to the surface during the filling process, and by
discharging the contents immediately into the cell at one corner,
the cover having been previously displaced at a slight obliquity to
admit the end of the pipette. With the filling of the cell the cover
automatically moves into place, and practice soon enables one to
fill the cell without inclusion of air bubbles. With the exception
of the heavier rhizopods, all of the organisms are as a rule very
evenly distributed by this method.
The identification and enumeration of the contents of the cell
were carried on with the help of a mechanical stage and a f Bausch
& Lomb objective, with a Zeiss C for higher magnification when
needed for the detection of fine details or for counting the smaller
organisms in the filter-paper catches.
After considerable experimenting, the following method was
established in the work of enumeration. Four sheets, each with
numbers 1 to 76 at the left, were fastened temporarily to accom-
panying key sheets, each number on each sheet standing for one of
the more common species. One sheet was assigned to algae,
diatoms, and miscellaneous organisms; and one each to Protozoa,
Rotifer a, and Entomostraca. As the plankton sample was examined
under the microscope the identifications were called off, and entered
on the sheets by a clerical assistant. Six of the most abundant
species were recorded by the observer himself on six tallying
machines registering 1,000, and conveniently arranged in a box at
his right. By adjusting the springs to give different sounds when
registry was made, and by modifying the surfaces pressed by the
fingers so as to differentiate the several machines without looking
at them, it w r as possible to use these without raising the eye from
the microscope, and thus to avoid the fatigue arising from the
repeated muscular readjustment of the eyes necessary when the
observer makes his own entries in a written record. Common
species not recorded by the tallying machines were generally abbre-
viated or designated by easily-called tokens. When once fairly
familiar with the species it was possible by means of these labor-
saving devices to make identifications and enumerations of several
heavy planktons per day.
By a number of tests I found that when the enumerations of a
species in a given collection reached 1,000, little was gained by
carrying it to higher numbers. A limit of error of 5 per cent.
can be thus obtained if the species in question is distributed evenly
in the cell and all precautions are observed to secure accuracy.
Enumerations were often carried beyond this point, but rarely beyond
3,000. The accessions numbers of the collections from our catalog
of collections served to designate each sheet of data and all note
slips bearing on the collection or its constituent organisms. When
the enumeration was completed, the factors of collection, dilution,
and enumeration were entered on the sheets, and the number of
individuals of all species represented was computed and carried to
the right of the sheet. The totals of the various groups for ex-
ample, diatoms or Cladocera were then added up and entered on
the sheets in differential colors. By the use of the key sheets the
number perm. 3 of water of any given species could be quickly ascer-
tained. Species not in the key were entered by name on the sheets.
When the enumeration of all collections was completed, the
numbers per m. 3 giving the seasonal distribution of the various
7
species and groups through the collections of 1894-1899 were drawn
up on uniform folio sheets, and the annual totals and averages com-
puted therefrom. With the data in these forms it is possible to
turn at once to the statistics of the plankton of a given day, or to
the seasonal distribution of any desired species.
ACKNOWLEDGMENTS.
I am indebted to Prof. S. A. Forbes, Messrs. E. B. Forbes, F. W.
Schacht, and R. W. Sharpe for many suggestions concerning the
Entomostraca; to Prof. Frank Smith for assistance with the Oligo-
ch&ta of the plankton ; and to Mr. A. Hempel for my introduction to
the Rotifera. The identification of the cosmopolitan species of the
fresh-water plankton of the Illinois River was greatly facilitated by
the most excellent library of the Illinois State Laboratory of Natu-
ral History, the accumulation of many years' careful selection by
its director, Prof. S. A. Forbes. The literature of fresh- water fauna,
and to a large extent of its flora also, is very fully represented therein.
The excellent Laboratory collection of identified Entomostraca
from European specialists was also of great service.
I am indebted to Mr. R. E. Richardson for valuable services as
clerical assistant, and for substantial help in organizing the great
mass of data resulting from the enumerations.
' Except as noted in the discussion in subsequent pages, I hold
myself responsible for all of the identifications of the species re-
corded. The enumeration is also all my own work, with the excep-
tion of that of the nauplii, of two species oi^ttifflugia, and of Pedi-
astrum in about one third of the collections, in which I had the
assistance of Mr. R. J. DeMotte, and that of the commoner Rotifera
in a few of the collections, which were counted by Mr. Richardson.
DEFINITIONS.
The term "plankton" was used by Hensen ('87) to designate
"Alles was im Wasser treibt." It was applied by him only to that
assemblage of marine organisms which float passively in the open
sea, without active recourse to shore or bottom, and unable by their
own efforts materially to change their location. The term has
since been extended also to assemblages of organisms in fresh water
which bear a similar relation to open water. This fresh-water
plankton has been designated in turn " limnoplankton " by Haeckel
('90), a word which in a restricted sense is retained for the plankton
of lakes, while that of rivers has been distinguished by Zacharias
('98a) as "potamoplankton," and that of ponds ('98) as "heleo-
plankton." These distinctions are based upon the nature of the
environing body of water, and the terms are convenient, though
the separation of these types everywhere in nature is difficult, if not
impossible. Owing to the smaller size of fresh-water basins as
compared with those of marine character, the shore and bottom be-
come more important as factors in the environment of the plankton.
Within the fresh-water environment we also find degrees of impor-
tance of the shore and bottom which in ascending scale dominate
in the lake, river, pond, and marsh. Although each of these repre-
sents distinct conceptions, in nature we find them imperceptibly
intergrading, and neither these conceptions, geographical nomen-
clature, nor local parlance give us any final criterion which will
enable us to use the terms with the precision which a scientific
terminology would demand. The distinctions between these forms
of fresh-water plankton must lie in the plankton itself, if anywhere.
As I shall attempt to show later, these distinctions, though appar-
ent, in some cases at least, are nevertheless of minor importance,
and depend very largely upon the relative predominance of the
adventitious littoral fauna and flora rather than upon distinctive
assemblages of eulimnetic species. The striking similarity of this
eulimnetic plankton in all these types of environment and in widely
separated continents is a biological phenomenon of far more sig-
nificance than these minor differences. These distinctions between
the different types of fresh-water plankton are thus more a matter
of terminology than of biological import.
Among the organisms found in open water there are varying
degrees of dependence upon the shore and bottom. Some, as
Cyclops and many of the lower algas, have life cycles in which no
encysted or quiescent resting stage has been found, and actively or
passively their whole existence is passed in the open water. They
are at all times components of the plankton; that is, are continuous
planktonts. Others, as Dinobryon, many of the Rotifera and Cladoc-
era, and, in fact, the greater part of the eulimnetic organisms, have
an encysted stage which as a winter egg or a cyst descends to the
bottom and remains there for a season. Such organisms only
periodically, wholly or in part, leave the open water for a littoral
or benthal existence. They are periodic planktonts. Some organ-
isms, such as many of the rhizopods and diatoms and Hydra, appear
in the plankton under certain conditions of temperature and food.
They temporarily adopt the limnetic mode of life as a result either
of a change in their specific gravity due to internal changes, such as
an increase of the gaseous or fatty contents of their protoplasm, or
to changes in the buoyancy of the water due to changes in
temperature or in substances in solution in the water, or because
of the abundance of food in the open- water. They become
under these conditions actively adventitious planktonts. Still other
organisms are released from their usual contact with or attach-
ment to the substratum, or from their association with debris
or vegetation of shore or bottom, by movements or disturbances in
the water, and are swept into the open water only to return again
to their customary habitat when conditions favor. Practically all
of the smaller organisms inhabiting the shore and bottom and the
debris and vegetation found thereon are liable thus to enter the
open water, and to be found in forced and temporary association
with the eulimnetic fauna and flora. They are passively adventi-
tious planktonts.
Another class of organisms which occur in the plankton are those
which either as internal or external parasites find in plankton organ-
isms either a host or a substratum for attachment. These are in a
certain sense passive planktonts, and they may be distinguished from
other passive planktonts as attached or parasitic planktonts. Sharp
lines between these various classes of organisms found in open water
can not be drawn upon distinctions based upon their degree of
dependence upon the bottom and shore. An equally vague line
separates the organisms of the plankton from those more active
forms which by virtue of their powers of locomotion are to a con-
siderable degree independent of waves and current, and are able
freely to maintain their position in their preferred habitat. Among
the organisms commonly included in the plankton, the flagellates,
rotifers, and Entomostraca exhibit some degree of activity, such as
is seen in their limited vertical migrations, while larger organisms,
such as Leptodora hyalina and the larvae of Corethra, are capable of
movement sufficient to give them considerable independence in the
matter of their position in the water. We thus find degrees of inde-
10
pendence which approach closely that found in young fish and the
large insect larvae organisms not always regarded as planktonts.
The plankton is thus a composite assemblage of organisms whose
association depends in varying degrees upon their relation to their
common habitat, the open water. In actual practice, all the organ-
isms found in the open water are regarded as within the scope of
plankton investigations, and justly so, for by virtue of their pres-
ence they become more or less involved in the complex interrela-
tions which pertain to the flux of matter, the succession of species,
and the food relations which exist through the changing seasons in
the aquatic environment.
In our own investigations it has been our purpose to include all
the organisms found in our collections ; that is, all which our meth-
ods of examination give us a sufficient means of investigating.
Naturally, the bacteria are to large extent excluded from our con-
sideration, though they properly belong to the plankton, and in the
processes of nitrification and denitrification play an exceedingly im-
portant part in the economy of aquatic life.
THE COMPOSITION OF THE PLANKTON.
The composite character of the plankton is especially marked in
streams , as, for example , in the Illinois River, owing to the mingling
of organisms from a great variety of tributary sources backwaters,
lakes, ponds, pools, marshes, swamps, brooks, rivers, canals, sewers,
drains, and industrial wastes. Few lakes possess so varied a supply,
and in none can the proportional effect of these contributions exceed
that of the stream. Added to this contributed assemblage, and in
some seasons predominating over it, is the indigenous or autono-
mous plankton of the stream itself.
The component organisms of the plankton of the Illinois River
number 528 forms, including only those which have been identified
from collections made in the main stream and including both
species and well-defined forms or varieties. Species found thus far
only in the backwaters are not included, though there is little doubt
that they occur also in the main stream. No effort has been made
to build up merely a long list of species, but only to identify, so far
as possible, the common and recurring forms. Neither has any
attempt been made to establish new species or revise those already
11
described, though a magnificent opportunity awaits the naturalist
who has the fortitude to analyze the exceedingly variable forms
which compose the plankton, and to determine by modern methods
which of these variants are entitled to specific rank. It has seemed
to the writer that the only satisfactory basis upon which species,
and pre-eminently those of the fresh-water plankton^ can rest, lies in
a careful determination of the limits of seasonal and local variation
within the area of distribution. This means breeding under con-
trol, and the study of variation by modern statistical methods.
Both of these lines of inquiry lie beyond the purpose of the present
paper, and plainly beyond the possibilities of accomplishment by
any one investigator, when the great number of species and the pres-
ent state of the literature of the subject is considered. It is becom-
ing constantly more evident that the species of the plankton are in
the main cosmopolites, and the world literature of the subject must
be taken into consideration in any thorough attempt to handle the
systematic side of the subject. During the progress of this work,
which was begun in 1894, every effort was made to secure all perti-
nent literature bearing on the genera of plants and animals repre-
sented in the plankton, and so far as possible in the enumeration of
the collections the individuals were referred to "species" already
described, or, in default of this, recorded as "unidentified." In
some groups notably the desmids, diatoms, and unicellular
algas it was not possible under the conditions of plankton enumer-
ation to apply to all the individuals enumerated the fine distinc-
tions which specialists in these groups have made. They have been
thrown under certain of the better-defined species, which thus stand
in our records as representatives of closely related variants as well
as of the types of the species named. Examples of this appear in
Closterium, where two species only were listed. Probably a num-
ber of so-called species among the scores described in this genus
will be found among the individuals in our plankton here referred
to the two species C. acerosum and C. lunula. So, also, in the case
of Melosira; two principal types were listed, M. varians and
M. granulata, though even these two seem at times to intergrade.
Other described species will be found among the individuals thus
distributed. In the case of Difflugia globulosa and D. lobostoma a
large number of intergrading and variable forms are included. It
would be possible to find among these, representatives of many
12
recently described species. In these instances the difficulty lies
not so much in finding representatives of these closely related
species, but, rather, in drawing the lines between them and placing
every individual enumerated in the proper pigeon-hole. To avoid
this difficulty, the separation was not attempted in every case.
With the hope that the results would throw some light on the ques-
tion of seasonal variation, this separation was attempted in the
genus Brachionus, where the species characters are confined to
prominent structural features.
So far as it was feasible, specific distinctions w r ere accepted as
found, and utilized whenever possible. In the lists and discussions
which follow, the inclusion of a species does not necessarily carry
with it the inference that it is regarded by the writer as valid or
well founded. It merely represents in our enumerations a more or
less continuous succession of organisms which conform approxi-
mately to the descriptions and figures of the species designated by
the name in question. Inferences regarding the rank or validity
of the species reported will be given whenever the statistical data
or my observations on the variability of the organism seem to
afford data bearing on the standing of the species. While not a
few of the species reported may justly be regarded as synonyms, an
effort has been made to use only names which represent valid
species or at least a variety or a seasonal form.
COMPARISON OF FRESH-WATER AND MARINE PLANKTON.
The plankton of fresh water is very generally composed of an
assemblage of organisms, of plants and animals, principally crypto-
gams and invertebrates. Not all orders are represented, and those
that do occur vary greatly in the number of their representatives.
The fresh-water plankton differs from that of the sea in the almost
universal absence of larval forms, in the smaller number of inverte-
brate groups represented, and in the smaller size of its component
organisms. Fresh-water plankton has almost no limnetic coelen-
terates, Hydra fusca being the only representative as yet discovered
in our locality. The absence of the larger Crustacea, of limnetic
mollusks and worms, and of tunicates and Radiolaria robs limnetic
life of the diversity found among pelagic organisms of the sea.
The only larval stages found in our locality are the glochidia of the
13
Unionidae, whose limnetic sojourn is at the best but brief, and the
larvas of certain dipterous insects, such as Chironomus and Corethra.
The limnetic habit of these larvas is hardly established as yet. The
small size of fresh-water planktonts as contrasted with those of the
sea is very striking. Representatives of the same group for exam-
ple, the Dinoflagellata and the Entomostraca in the two habitats
exhibit this contrast. The largest entomostracan of fresh water is
less than a centimeter in length, and there is nothing to compare
with the pelagic ccelenterates, Mollusca, or such tunicates as Salpa
and Pyrosoma. The smaller size of fresh- water planktonts may
be due to the lower specific gravity of the environing medium, and
perhaps also to the effect of smaller quantities of dissolved salts
upon the metabolic processes of limnetic animals.
Notwithstanding this absence of large individuals in the plank-
ton of fresh water, the total quantitative production of plankton
per cubic meter is greater here than in the sea. For example, the
average production in the Illinois River is 2.71 cm. 3 , and the average
amount in adjacent backwaters rises as high as 22.55 cm. 3 (in Phelps
Lake) . These measurements were made by the centrifuge, and the
results of the "Plankton Expedition" of Hensen reduced to this
basis of measurement by Kramer ('97) show that the Atlantic
Ocean at the time of this expedition had in the upper strata exam-
ined but 0.12 to 0.48 cm. 3 of plankton per cubic meter
ORGANISMS OF THE PLANKTON.
The groups of plants represented in the plankton of the Illinois
River are principally algas, of which the Bacteriacea are but partially
retained in the collections and are usually omitted in plankton
investigations. The Sckizophyce&, or blue-green algae, furnish a
few important representatives and a number of adventitious .species.
The ChlorophycecB, or green algas, on the other hand, abound both
in species and individuals, and afford an element of great impor-
tance in the primal food supply. The Bacillariacece are exceedingly
abundant, and are represented by a number of eulimnetic, as
well as many adventitious, species. They also constitute one of
the primal sources of food for the zooplankton. The Conjugate
furnish but few species and individuals -principally desmids to
the phytoplankton. The phanerogams afford a few species which
14
are often taken with the plankton by virtue of their semi-limnetic
habit, but do not in the living state enter the food cycle of the
plankton nor affect its economy except as competitors.
The zooplankton includes representatives of a considerable
range of groups, though both in species and individuals the Proto-
zoa, Rotifer a, and Entomostraca predominate among the animals.
Representatives of other groups are in the main adventitious.
Among the Protozoa, the Rhizopoda are constantly represented
by many individuals and a considerable number of species, many
of which may be adventitious, but most of which are wont to adopt
the limnetic habit during the warmer months. The Heliozoa are
few both in species and individuals. The Mastigophora (which in
our discussions include all green and brown flagellates often clas-
sified with the Chlorophyce and Ph&ophycece] vie with the Chloro-
phycecs and Bacillariacecz for the first place as converters of the
inorganic (and perhaps also the dissolved organic) matter into food
for the zooplankton. They are exceedingly numerous in our plank-
ton both in species and individuals, and form quantitatively a con-
siderable part of the plankton during the summer months. The
usual method of plankton collection by silk bolting-cloth per-
mits a large proportion of these organisms to escape. The Ciliata
furnish a few constant members of the plankton, and numerous
adventitious and parasitic species. During the low water of
autumn, when bacterial contamination is at its height, these organ-
isms form a large part of the plankton. The small size of some of
the ciliates, combined with their motility and flexibility, renders
the loss by their escape through the silk net considerable. The
Suctoria furnish but few species and individuals mainly adventi-
tious or attached to other planktonts.
The Rotifera constitute, both in species and .individuals, the
most important single group of analytic organisms, that is those
of distinctly animal metabolism, occurring in our plankton. This
may in part be due to our shallow warm waters and to the abundance
of Chlorophyceoz and Mastigophora, which enter largely into their
food. This abundance of the Rotifer a may prove to be character-
istic of the plankton of rivers ( potamoplankton) as contrasted
with that of lakes (limnoplankton) . While many rotifers are
eulimnetic, the plankton also contains numerous adventitious
species.
15
The Entomostraca include the largest fresh-water planktonts,
and in every respect constitute an important element of our river
plankton. They form the final link in the food cycle which con-
nects the nutrients in solution in the water and in decaying detritus
with the fish and other aquatic vertebrates. They include numer-
ous species, some of which are adventitious. All of the Ostracoda
belong to this latter class. The Cladocera furnish some of the
most important eulimnetic species and a large number of adventi-
tious forms, while the Copepoda are almost wholly eulimnetic.
In addition to these groups, the Turbellaria, Oligoch&ta, Hexap-
oda, Hydrachnida, Gastrotr-icha, and Bryozoa furnish a few species
and individuals of a semi-limnetic or adventitious character to the
plankton.
In the table which follows, these various groups are listed, and
the number of forms occurring in each is noted. In order to give
some idea of the proportionate representation of these groups in
our plankton, the table includes the sum of the number of indi-
viduals per m. 3 of water in the weekly collections for the year 1898.
This was a year of no marked departure from the normal regimen
of hydrographic conditions (Part I., PI. XII.). The summer and
autumn flushes tend to lower the population somewhat below that
of more stable seasons, but beyond this feature there is nothing to
suggest that the plankton of this year may not represent a fair
average of that recurring each year in the Illinois River. The fig-
ures given, in all cases refer to the number of individuals per cubic
meter (excepting only such cases as Synura and Uroglena, where
the colony rather than the individual becomes the unit). The algae
and Protozoa include many species enumerated in filter-paper col-
lections, which accounts for the large numbers in some of the totals.
The "number of forms" listed refers to the total number found in
the waters of the river during the period of our operations. Some
species not noted in 1898 are therefore included. Unidentified
forms are not included in the list of number of species, though the
groups here listed to which they belong were known. Some forms
referred to genera but not determined as to species are, however,
included.
This table throws some light upon the ecological relations of
the groups composing the plankton, since it gives some clue to their
relative numbers, and these condition in a general way the food
16
relations existing between the different groups. The plants are
more abundant (and generally smaller) than the animals, outnum-
bering them nearly 5 to 1 . Computation shows that for each one of
the Cladocera there are 7 Copepoda, the predominance of the latter
CONSTITUENT GROUPS OF THE ANNUAL PLANKTON OF THE ILLINOIS RIVER.
AVERAGE OF 52 WEEKLY COLLECTIONS IN 1898 NUMBER PER M 3 .
Number of
forms
recorded.
Number of
individuals.
Algae:
Bacteriaceae
3
(57,142,822)*
Schizophyceae
9
85,909,985
Chlorophyceae
33
53,175,105
Bacillariacece
29
396,192,716
Conjugatas
7
48,459
Phanerogamia
2
9
Total phytoplanktonts
83
535,326,274
Protozoa total
(185)
(111,731,000)
Mastigophora
68
95,856,449
Rhizopoda
59
55,364
Heliozoa
5
4,871
Sporozoa
3
1,638
Ciliata
45
15,812,346
Suctoria
5
332
Rotifera
104
592,416
Entomostraca total
(43)
(47,041)
Cladocera
26
6,242
Ostracoda
4
191
Copepoda
13
40 , 608
Miscellaneous . .
114
9,393
Total zooplanktonts
446
112,379,850
Total planktonts enumerated.
529
647 706 124
Synthetic (chlorophyll-bearing)
613 017,986
Analytic (non-chlorophyll-bearing)
34 687 781
being accounted for in part by the fact that their larval stages are
free-s Dimming and appear in the enumerations, while the young of
the Clidocera are not set free until nearer maturity. About 10 to
20 per cent, of the Copepoda are adults. The relative numbers of
* Represents fragments of filaments, and is not included in totals.
17
the two groups are not so disproportionate as the figures might
seem to indicate. For each one of the Cladocera there are 95 roti-
fers and almost 18,000 Protozoa. The latter are distributed as fol-
lows: There are 9 rhizopods, almost 2,400 ciliates, and over
15,000 flagellates for each one of the Cladocera. There are also
about 86,000 plants for each of these Cladocera. Of these plants,
64,000 are diatoms, 14,000 are Schizophycecz, 9,000 Chlorophycece,
while but 8 are desmids. The great abundance of diatoms, of
green and blue-green algae, and of chloryphyll-bearing flagellates
affords, it would seem, an abundant food supply for the zooplankton.
If of the Mastigophora the colorless flagellates only be retained in
the zooplankton, and the remainder which are predominantly
synthetic forms be included with the phytoplankton, we find the
latter outnumbering the analytic organisms (zooplankton) 18 to 1.
Quantitative values in the matter of food relationships are not
readily determined except by a combination of the chemical and
experimental method. These results by the statistical method
express, with more or less error, the equilibrium of the biological
components in terms of the individual organisms.
DISCUSSION OF THE STATISTICAL DATA OF THE SPECIES COMPOSING
THE PLANKTON OF THE ILLINOIS RIVER IN 1894-1899.
In the following pages the organisms occurring in the plankton
of the Illinois River will be recorded, and from the statistical
data accumulated by the enumeration method, facts pertaining to
their relative abundance, seasonal distribution, and periods of max-
imum occurrence will be cited. The average number per cubic
meter for the year 1898 will be given, based upon the averages of
52 collections distributed regularly throughout the year (Part I.,
Table III.). This year is chosen because of the regularity of the
times of collection and the absence of any considerable irregularity
in the hydrograph. Statements concerning seasonal distribution,
etc., are based upon the records for all the years 1894-1899. All
figures pertaining to species or groups marked with an asterisk,
and starred figures elsewhere, are based upon filter-paper catches;
all others, upon those of the silk net. Temperatures are in Fahren-
heit, and are of surface waters at time of collection..
The margin of error in statistical work of this sort is confessedly
large. The complex character of the data with which I am deal-
ing, and especially the extreme range in numbers, have made it
necessary that I should adopt some consistent method of treating
the computations. I have therefore chosen to carry out the num-
bers to units, as the most feasible method of avoiding confusion in
the handling of the data. The use of round numbers would have
been just as accurate. Computation to units is therefore to be
understood as a matter of convenience, and not as an effort to
exhibit a false and unattainable accuracy.
CRYPTOGAMIA.
BACTERIACE^;.*
Records were kept of the masses of the larger members of this
group which occurred in our plankton catches. They were princi-
pally the dichotomously branched brownish fragments of Creno-
thrix, filaments of Beggiatoa, and colonies of Micro coccus. The
average number recorded for this year was 57,142,822, and they
occur throughout the year in every collection, rarely falling below
18
19
10,000,000 per m. 3 , and reach their maximum development (over
600,000,000) in winter months (December to February), especially
during low water and more stable conditions, as in January, Feb-
ruary, and December, 1898 (Pt. I., PL XII.). At such times the
temperature is at or near 32. With flood conditions and rise in
temperature the numbers fall below 100,000,000, Tunning from
10,000,000 to 50,000,000 during most of the summer. The decline
is due in part to the dilution by flood waters, and largely to the
retreat up the stream of the crest of the wave of bacterial activity
caused by the Peoria pulse of sewage. As noted in the discussion
of the chemical conditions, in Part I., this wave lies considerably
above Havana during the warmer months. Summer floods, as in
June and September, 1897, are wont to wash into the river large
quantities of these organisms, bringing the numbers up to 300,-
000,000 at times. The figures above cited give but a feeble repre-
sentation of the real conditions in the river during this period of
maximum. Many of these organisms become attached to objects
along shore, and accumulate in great quantity in quieter waters
along the channel. They form a serious menace to the fishing
industry, since they accumulate in a day or two upon the fyke-nets
in quantity so great that their- weight and resistance to the current
are sufficient to break down the nets. Their effect upon the consti-
tution of the plankton is seen in the marked increase in certain
ciliates which accompanies the maximum of these organisms.
SCHIZOPHYCEyE.
Nine forms were recorded, though a number of others which
occurred but rarely in the plankton remained unidentified. The
average number (combined silk and filter-paper records, but omit-
ting the former when the latter are available) is 85,909,985 per m. 3
This group contributes to the plankton throughout the year, and
though numerically abundant is quantitatively less important,
owing to the small size of its most abundant member, Microcystis.
This species and Oscillatoria constitute quantitatively the greater
part of the blue-green algas of the plankton. In contrast with the
plankton of Lake Michigan, there is a noticeable decrease in the
proportion of AnabcBua and Clathrocystis. Rivularia, Gloiotrichia,
and Aphanizomenon flos-aquce, often reported in fresh- water plank-
(3)
20
ton, were not found in our fluviatile environment. This group
contributes to the water-bloom, contains a number of adventitious
planktonts, and is one of the primal sources of the food supply.
In our waters it seems to be quantitatively much less important
than either the Chlorophycea, the BacillariacecE, or the synthetic
Mastigophora.
DISCUSSION OF SPECIES OF SCHIZOPHYCE^E.
Anabozna spiroides Klebahn.* Average number, 637,692 (silk
15,431). In the water-bloom from the last of June till the end of
October. Not noted in 1898, but not infrequent in 1897 a low-
water year. Temperature range, 60-89. Data insufficient to
determine maximum. Largest number recorded, 7,200,000, June
28.
Clathrocystis ozruginosa (Kutz.) Henfr. Average number of
colonies or masses, 83. More abundant in the previous low- water
year. From May till the end of November in the water-bloom.
Predominantly a midsummer species. Maximum in August and
September (108,000) . Confined principally to the low water of mid-
summer, appearing when the water reaches a temperature of 70,
and reaching its maximum development in temperatures above
this point, declining at once to small numbers (less than 1,000)
when the temperature falls below 60, but lingering till the water
approaches the freezing point late in November.
Merismopedia glauca (Ehrbg.) Nag. Average number of col-
onies, 93. In 1897, 889,412.* In the water-bloom. Recorded
from July till the end of October, and also singly in January and
February. It was more abundant in 1897 than in 1898, and the
maximum number (15,840,000*) appeared on August 31.
Microcystis ichthyoblabe Kutz.* Average number, 83,059,615.
Recorded in all collections throughout the year, except in some
flood waters of February and March, when the silt probably ob-
scures it. Minimum numbers (less than 50,000,000) prevail during
cold months, November to April, when the temperature ranges
from 32 to 50. A well-sustained pulse exceeding 200,000,000
appears with the volumetric plankton maximum of April-May
(Pt. I., PI. XII.) and declines to the previous minimum with the
falling off in the plankton. The maximum pulse appears later, in
August and September in 1898, in September and October in 1897,
21
averaging about 200,000,000, and reaching 1,697,000,000 August 9,
1898. The temperatures during these pulses are above 60, and
the period of the maximum comes toward the close of that of max-
imum summer temperatures, and sometimes in the autumn decline
(Pt. I., PL XI. and XII.), when low and often stable jriver-levels
usually prevail. A vernal and an early autumnal pulse are thus
both present in the distribution of this species. It is not improb-
able that other species than the one named have been included in
the enumeration along with it on account of the small size and lack
of striking characteristics. There are suggestions of recurrent
pulses at intervals of 2-6 weeks in the records (Table I.).
Oscillatoria spp. Average number, 15,431 (filter-paper, 637,692).
The probable inclusion of several species in the sums under this
heading may account in part for the irregularity of the seasonal
curve. Oscillatoria has appeared in every month of the year,
though the occurrences were most frequent in the period from July
till the first of October. The numbers are exceedingly irregular
and variable, and the pulses of numbers seem to attend the initial
stage of floods following stable conditions. Thus, while these
organisms occurred but singly or sparingly in the plankton during
the autumn of 1897, they rose to 277,200 with the flood of January
11, 1898, doubtless torn loose by the current from the bottom-
their normal habitat. They are thus usually adventitious addi-
tions to the plankton. Their frequent irruption into the plankton
during midsummer and early autumn, and to some extent at other
times, is due in part to the evolution of marsh gas in the detritus on
the bottom. This breaks up the mats of Oscillatoria which coat
the bottom and distributes them through the upper levels, where
they remain in suspension for some time. This phenomenon is
more prevalent in the marshy backwaters than it is in the river.
Flood invasion in midsummer into the backwaters, such as Quiver
Lake, is wont to cause there stagnation and great increase in Oscil-
latoria, which to some extent enters the river with the run-off of the
flood. Movements in the water and the evolution of marsh gas are
thus principally responsible for the presence of Oscillatoria in the
plankton. It still remains possible that its flotation during periods
of optimum conditions of growth may be due to internal physio-
logical conditions which lower the specific gravity of the organism.
Its great abundance at times in upper levels in the backwaters sug-
22
gests the action of this factor, and if this be true, it becomes a tem-
porary rather than an adventitious planktont. Temperatures seem
to bear little relation to the occurrence of Oscillatoria in the plankton.
Tetrapedia emarginata Schrod.* Average number, 242,308.
From the first of August till the end of October in numbers from
1,000,000 to 3,500,000 per m. 3 , appearing later and in larger num-
bers in October in 1897 than in 1898. At temperatures above 65.
Tetrapedia gothica Reinsch, Glceocapsa polydermatica Kutz., and
Gloeocapsa sp. were recorded once or twice in the midsummer plank-
ton in relatively small numbers.
CHLOROPHYCEyE.
(Plates I. and II.)
Average number, 53,175,105, including, without duplication,
species from both silk and filter-paper collections. In 1897 this
was very much greater (139,739,850), owing to the prolonged low
water and higher temperatures of the late autumn. Although
abundant, these organisms are outnumbered by the diatoms six to
one, and by the synthetic Mastigophora by about two to one. The
ChlorophycecB of the plankton, with few exceptions, are minute, and
generally escape through the silk net. Pediastrum and colonies of
Botryococcus are about the only species of which the usual method
of plankton collection in our waters affords a fair representation.
The Chlorophycea appear in every collection examined through-
out all the years of our operations, with the exception of eight in
midwinter floods in 1895 and 1896. As a group they are adapted to
the whole range of temperatures, and exhibit in 1897, on April 28,
a well-defined vernal pulse of 367,200,000, and a series of autumnal
pulses culminating September 21 at 216,000,000, October 19 at
367,200,200, and November 23 at 52,000,000. In this year the
-midsummer pulses are of minor importance in comparison with
those of spring and autumn. In 1898 the vernal pulse is also well
defined, culminating May 3 at 212,406,400, and it is followed by a
series of four midsummer pulses of considerable magnitude, w r hich
culminate June 14 at 46,000,000, July 19 at 277,000,000, August 9
at 370,000,000, and August 30 at 189,000,000. The autumnal pulse
appears September 27, attaining 70,526,400. The summer and
autumn hydrographs of this year are much more disturbed than in
23
the previous year (cf. PL XI. and XII., Pt. I.), especially at the time
of the autumnal pulse. This may account for the contrast in the
two years. The Chlorophycea as a whole exhibit (PI. I. and II. and
Table I.) the tendency to form a seasonal curve of recurrent pulses
at approximately monthly intervals (three to six weeks)-which gen-
erally coincide with those of other chlorophyll-bearing organisms.
Thirty-three forms of Chlorophycecz were recorded, and closer
inspection of the collections will undoubtedly yield a considerable
additional number either of closely related, and therefore included,
species, or of those which occur but occasionally or in small numbers
in the plankton.
Numerically the leading species in the order of their importance
are Scenedesmus quadricauda, Crucigenia rectangularis, Actinastrum
hantzschii, Raphidium polymorphism, Scenedesmus genuinus, S. obli-
quus, Richteriella botryoides, Ophiocytium capitatum, Oocystis naegelii,
Ccelastrum cambricum, Oocystis solitaria, and Schroederia setigera.
With the exception of Botryococcus braunii and the species of Pedias-
trum, the remaining forms are both quantitatively and numerically
of minor importance. The species just named were enumerated only
in the silk-net collections, and ccenobia rather than individual cells
were listed. If allowance is made for the loss of small individuals
through the silk, and for the increase that would follow if individ-
uals rather than ccenobia were the basis of representation, Pedi-
astrum would occupy a place in the front rank of importance in the
ChlorophycecB of the plankton numerically as well as quantitatively.
As quantitative factors in the ecology of the plankton, Pediastrum,
Scenedesmus, Ccdastrum, and Botryococcus take precedence over the
smaller, though more numerous, forms, such as Raphidium and
Crucigenia.
The group is thus well represented in our plankton both in
species and individuals. The leading planktonts of the group
reported in European and other waters in lakes and rivers are here
represented almost without exception by identical or closely related
species. Botryococcus alone seems to be less abundant than in
lakes at least, according to my own observations, it is. much more
abundant in the summer plankton of Lake Michigan than in that of
the Illinois River. The maximum numbers of Pediastrum reported
by Apstein ('96) for Dobersdorfer See in July, when reduced to
number per m. 3 , are frequently equaled or surpassed in our waters.
24
Data for comparisons in the case of the more minute organisms
which escape the silk are lacking, since results of supplementary
methods have not, up to the present, been published elsewhere.
It seems probable, however, that the Chlorophycecs will be found to
be somewhat more characteristic of the plankton of rivers than of
lakes, and to be more prevalent wherever the shore with its decay-
ing vegetation forms a large factor in the environment or where
sewage contamination affords the requisite food for their develop-
ment.
DISCUSSION OF SPECIES OF CHLOROPHYCE^E.
Actinastrum hantzschii Lagerh.* Average number, 199,038
(silk net, 338). From May until the middle of November, with
maximum of 21,600,000 on August 30, 1898, and of 122,000,000 on
September 21, 1897. There are also indications of a vernal pulse,
which on May 25, 1897, attained 90,000,000. The major pulse
occurs late in the summer, in August and September, while dimin-
ished numbers continue until the first of November. Three single
occurrences were noted in January, 1898, following the unusual
prevalence of 1897, but aside from these the species occurs in the
plankton at temperatures above 45, and both pulses lie in temper-
atures above 65. As in many other species, a greater development
was attained in 1897, in stable low water, than in 1898 in disturbed
hydrographic conditions. This species occurs in the water-bloom, is
favored by stable conditions, and finds its optimum temperature
between 65 and 80.
Botryococcus braunii Kiitz. Average number of colonies, 75. In
previous years it was much more abundant, averaging 3,300 in 1897.
It occurs from the first of April well into October, though in 1897 it
continued until the middle of December. It may thus appear
throughout the whole range of temperatures, 32 to 90, but as a
rule occurs above 60. There is a suggestion of a minor pulse in
June, 1896, but not in other years. The major pulse attains
57,200 on August 15, 1896, and 42,000 on September 14, 1897, and
appears, with smaller numbers, in August of preceding years. The
species occurred but sparingly in 1898. It is found in the w T ater-
bloom, and is more abundant in the backwaters than in the main
stream.
Ccelastrum cambricum W. Archer.* Average number of cceno-
bia, 640,384 (silk, 477). Occurs from the latter part of March till
25
towards the end of November, but principally from May through
October. There are but slight indications of a vernal pulse, which
on May 25, 1897, culminates at 3,600,000. The major pulse cul-
minates at 10,800,000 on August 9, 1898. In the low water and
prolonged high temperatures of 1897 the major pulse continues
through September, culminating on the 21st at 32,000,000. The
average number in this year was about four times as great as in
1898. The temperature limit is 43, though occurrences are few
and numbers small below 65. The maximum development appears
within the period of maximum heat, and towards its close. It is
characteristic of the plankton of late summer and early autumn.
Crucigenia rectangularis Nag.* Average number of colonies,
7,153,846. Recorded in all months but March and April, but spar-
ingly from November till May. In 1897 pulses appeared in August,
September, and October, attaining 32,400,000, 57,600,000, and
118,800,000, respectively. In 1898 there was but a single pulse
in August, of 158,400,000. It was more abundant in the former
year. It is present continuously in large numbers from July to
October, though in 1897 the impetus of the unusual development
was manifested by the continuance of the species even into Janu-
ary. The optimum temperatures lie above 70, in the latter part
of the period of maximum heat, though the species has been found
in the plankton throughout the whole range of temperatures. The
abrupt decline in numbers occurs between 65 and 40. It is char-
acteristic of the plankton of late summer and early autumn.
Golenkinia radiata Chodat. Average number of colonies,
519,231. It appears most abundantly during the April-May plank-
ton pulse (7,200,000) and again, in increased numbers, at the end of
August, thus suggesting a vernal and a late summer maximum. It
seems to be most abundant at about 60, a temperature somewhat
below the optimum for the two preceding species. Two occur-
rences in December, 1896, and large numbers in August indicate its
adaptability to the full range of temperatures.
Oocystis naegelii A. Br.* Average number, 207,692. In 1897,
much more abundant (average, 4,243,235). Present in numbers
(over 5,000,000) from the end of May till the end of September.
In 1897, pulses of 10,800,000, 46,800,000, and 24,750,000 appear in
May, July, and September respectively. Both numbers and oc-
currences are much less in 1898. The optimum conditions thus lie
26
above 70, though isolated occurrences in March and December in-
dicate its presence throughout the whole range of temperatures.
It appears to be a summer planktont without the marked prefer-
ence for the close of the period of maximum heat noted in some
other ChlorophycecE.
Oocystis solitaria Wittr.* Average number, 121,153. In 1897
much more abundant, averaging 2,170,588. In this year it
occurs in numbers above 1,000,000 from the end of July till the end
of October, reaching a maximum of 36,000,000 on September 21,
1897. Its optimum conditions occur during the latter part of the
period of maximum heat, at temperatures approaching 80. It
disappears at 60, save for isolated appearances in December, at
33- - a fact which suggests its persistence in small numbers
thoughout the year. It is characteristic of the plankton of late
summer, that is, of low water, high temperatures, and stable con-
ditions.
Ophiocytium capitatum Wolle*. Average number, 1,465,385.
More abundant in 1897, averaging 2,858,823. Present from the
last of April until the beginning of November. There is some indi-
cation of a vernal pulse, which on May 25, 1897, attains 3,600,000,
and on April 26, 1898, 10,800,000. The major pulse appears in
late summer or early autumn, attaining 57,600,000 on September 21,
1897, and 28,800,000 on August 9, 1898. The two pulses are
separated by an interval in which occurrences are less frequent and
numbers smaller. This planktont thus exhibits the tendency
towards seasonal maxima near the average temperature. The
greater development in 1897 is followed by a prolongation of the
occurrences into November. The optimum temperature appears
to be about 60 or above, the vernal pulse appearing at that tem-
perature, and the major one at 71. No records occur below 46.
Pediastrum boryanum (Turp.) Menegh. Average number, 4,510.
This alga was found in every month of the year, though not in
every collection examined. The numbers present fluctuate greatly
and are usually much less than those of P. pertusum, with which.it is
associated, and with which it fluctuates, often with remarkable
coincidence. I have included under this head those individuals
in which the ccenobium is a plate with no intercellular spaces or
only insignificant ones. Individuals are not lacking which serve
to connect this species with P. pertusum, and, indeed, with others
27
which have been described in this genus. This genus includes the
most abundant of the larger algae in the plankton of fresh waters,
and it affords an attractive field for the study of variation by statis-
tical methods and for the determination by the experimental method
of the effect of environmental changes upon structure. The two
groups of individuals included here under P. boryanum and P.
pertusum give typical curves of seasonal distribution which are so
similar that their combination in a single series would not greatly
modify the resultant seasonal curve. In the sum total of all
collections P. boryanum (1,034,000) includes about one tenth of the
number referred to P. pertusum (10,830,117).
A few scattering individuals, generally less than 1,000 per m 3 .,
appear at irregular intervals, during the colder months, from the
first of December until the end of March. The number increases as
the temperature rises, and the species appears in all collections un-
til November, when it again becomes irregular in its occurrence in
the plankton. The fluctuations in numbers during this period are
very marked, the pulses of frequency being set off by intervals in
which the numbers are small. A slight pulse of 2,120 appears on
November 17, 1894. In 1895 the vernal pulse attains the very
unusual number of 572,824 in the unusually low water of that year,
and the autumnal pulse of September 5 is but 10,600, and is followed
by a secondary one on November 27 of 4,081, perhaps as a result
of the stable conditions and the abnormally high temperatures
(above 45) which then prevailed (Pt. I., PI. IX). In 1896 the
vernal pulse culminates May 18 at 31,164, while the autumnal pulse
is scarcely visible and the numbers throughout the summer are
small, as a result, it may be, of the repeated floods of that year
(Pt. L, PI. X). In 1897, with few vernal data, the vernal pulse does
not appear, though a rise to 8,000 occurs on July 21. The major
autumnal pulse culminates on September 14 at 14,400, and another
one on October 12 at 6,000, attending the late autumn of that
year. In 1898 there are vernal pulses on May 10 of 6,400 and on
June 14 of 32,000. The autumnal pulse on September 27 reaches
the considerable number of 65,600. In the winter of 1898-99
Pediastrum was seemingly absent from the plankton. The pulses
are thus somewhat irregular, though there is in this species a
suggestion of vernal and autumnal pulses at corresponding
28
temperatures. The optimum conditions seem to lie above 60
and the maximum numbers to occur at or near 70.
Pediastrum pertusum Kiitz. Average number of coenobia,
44,372. This species appears in the plankton in all months of the
year and in almost all of our collections. It is the most abund-
ant representative of the Cklorophyceaz which is retained by the
silk of the plankton net, and is quantitatively an important
factor in the ecology of the plankton. The numbers during the
colder months, from November to April, when the water is from
32 to 40, are few, and the sequence of their appearance is fre-
quently interrupted. As the temperature rises in April the num-
bers increase, and the vernal pulse culminates in a maximum in May
or June. There is no indication of the vernal pulse in the scattered
collections of 1894. In 1895 the pulse is extreme, reaching 5,264,860
on June 19, in a period of exceptionally low water. In 1896 a pre-
liminary vernal pulse culminates May 8 at 23,580 and is followed on
June 17 by one of 107,200. In 1897 the few spring collections do
not reveal any vernal pulse, while in 1898 a minor one on May 17
reaches 5,600, declines to 600 at the end of the month, and rises
again to 56,000 by June 21. These vernal maxima all occur or at
least pass through their period of development before the water
reaches its midsummer temperature of approximately 80. They
develop during the transition from 60to 80 (Pt. I., PI. IX. to XL).
Autumnal pulses during the decline from 80 to 60 appear on Sep-
tember 5, 1895, (105,996), on September 30, 1896 (9,200), on Octo-
ber 12, 1897 (231,200), and on September 27, 1898 (259,200). In
addition to these pulses there are others at irregular intervals during
the summer: on July 30, 1894 (154,548), on July 2, 1896 (68,400),
on August 15, 1896 (22,000), on July 14 (289,600) and on August
31, 1897 (442,000), and on August 2 (295,200) and 30 (326,400),
1898.
The optimum conditions of development thus lie above 60,
and pulses are more frequent in spring and late summer or early
autumn near 70, though they appear somewhat less frequently
during the summer in our maximum temperatures near 80. The
cause of these pulses is not conclusively demonstrable from the data
at hand, owing in part to the interval between examinations.
Daily examinations of the plankton and chemical analyses seem to
be desirable for such demonstration. There are indications, how-
29
ever, that certain conditions in the environment increase the
amplitude of the pulses by hastening the rapidity of reproduction of
these organisms. Of the fifteen well-defined pulses appearing in
our records of six years, all but three minor ones occur in stable con-
ditions, such as pertain to sustained low water. The ""greater part
of these pulses, however, occur in declining floods, when contribu-
tions from backwaters are considerable. It may seem ill-advised to
refer to the conditions of falling river-levels as "stable"; neverthe-
less, they are relatively much more stable than those which attend
the in-rush of silt-laden flood- waters, and involve fewer changes in
factors of the environment. Save in the matter of the relative con-
tributions of backwaters and of sewage dilution they resemble
those of sustained low water. These Pediastrum pulses are also re-
lated to the nitrate pulses (Pt. I., PI. XLIII.-XLV. and Table X.),
but the relation is not uniform. In the majority of instances the
pulses of 1896-1898 (during which time chemical anaylses are
available) coincide approximately with the crest or decline of in-
crease in nitrates. For example, the pulse noted on July 17, 1896,
of 107,200 from a previous level of 1,210 on June 1, follows a wave
of nitrates 'progressing for three weeks and culminating on June 9 at
3.25 parts per million arise from 1.5 (Pt. I., PI. XLIIL). On June
16 the nitrates have fallen again to 2.2, and on the 23d to 2.0, but
rise on the 30th to 2.8. Pediastrum responds to these changes by
dropping from 107,200 on the 17th to 15,000 on the 27th, and by
rising again on July 2 to 68,400. Not all of the fluctuations in the
two are concomitant. Some of the most marked pulses of Pedi-
astrum appear at the lowest levels of the nitrates. For example,
that of August 30, 1898, of 326,400, follows no nitrate wave, though
it coincides with a reduction in nitrates to the minimum of .05. On
the other hand, the nitrites had just passed on August 23, an un-
usual pulse, to .42, falling again on August 30 to .22 and on Septem-
ber 6 to .05 with the passing of the Pediastrum pulse. Pulses of
Pediastrum are thus apparently not dependent for their develop-
ment upon an abundance of nitrates above the levels shown in the
analyses, though a decline in these sources of food or in other forms
of nitrogen usually attends these pulses. Pediastrum is but one of
many factors among the planktonts, and in the environment,
biological and chemical, concerned in these changes, and con-
clusive demonstration of its ecological relations must be obtained
30
by the experimental method. The data here cited are suggestive
only; not conclusive.
The relation of Pediastrum to the volumetric pulses of the plank-
ton is not a constant one, though there is some correspondence in
their fluctuations. The extreme maximum (3,264,800) of June 19,
1895, is coincident with a plankton pulse of 30.42 cm. 3 , but the num-
ber of collections is insufficient to show the relative fluctuations of
the plankton and Pediastrum at that season. In May and June,
1897, and in October, 1898, the Pediastrum pulses culminate shortly
after the volumetric pulses. In July and September, 1897, arid
in August, 1898, they coincide.
Polyedrium trigonum Nag.* Average number, 432,692. Ap-
pears from June through September, disappearing when falling
temperatures reach 60. In 1897 it continues through October
with the higher temperatures (averaging 65) of that year. There
are slight indications of a September pulse.
Polyedrium trigonum forma minus Reinsch and var. tetragonum
(Nag.) Rabh., P. bifurcatum Wille, and P. gracile Reinsch, were
also recorded in a few collections during the period of occurrence
of P. trigonum. They are all evidently summer planktonts.
Raphidium polymorphum Fresen.* Average number, 2 1 ,450,000.
Occurs in every month of the year and in a majority of the collec-
tions. In 1897 a vernal maximum of 201,600,000 occurs on April
27 and an autumnal one of 28,800,000 on September 21. In 1898
a vernal pulse culminates May 3 at 24,000,000, and thereafter
throughout the summer at intervals of three to six weeks there
occur five other pulses, the greatest of which culminates July 19 at
75 ,600,000. A pulse of 90,000,000 on a declining flood in February,
1899, indicates an adaptation on the part of this organism to the
whole range of temperatures. A pulse of 25,200,000 December 3,
1896, further illustrates this adaptability. Records in 1897 and
1898, however, suggest that the optimum lies above 60. It is thus
a perennial planktont.
Raphidium longissimum B. Schroder. Appeared sparingly in
February, August, October, and December, suggesting that it has
also a perennial distribution.
Richteriella botryoides (Schmidle) Lemm.* Average num-
ber, 6,399,705 (in 1897). From May to November, with a vernal
pulse of 25,200,000 on May 25, and an autumnal one of 100,800,000
31
on September 21. Optimum temperature about 70, and disappear-
ing from our records below 60.
Scenedesmus bijugatus (Turp.) Kiitz.* Average number, 155,769.
Sparingly from May till the close of September, with slight traces of
vernal and autumnal pulses.
Scenedesmus denticulatus Lagerh.* Average number, 86,538.
A few occurrences in late summer and early autumn.
Scenedesmus genuimts Kirchner.* Average number, 778,846.
From May till the first of October, but continued through this
month in 1897. Vernal pulse not observed, though the autumnal
pulse attains 28,800,000 on September 21 and October 26, 1897.
Midsummer pulses appear in 1897 on July 14 (16,200,000), August
17 (14,400,000), and in 1898 on August 9 (19,800,000). Optimum
temperatures lie above 60, though an occurrence in December in-
dicates the adaptability of this organism to lower temperatures.
Scenedesmus obliquus (Turp.) Kiitz.* Average number, 1 ,505 ,769
(silk, 673). This form appears in our records from the last of April
until the middle of November. Traces of vernal and autumnal
pulses appear in both 1897 and 1898, with intervening midsummer
fluctuations of even greater magnitude. In 1897 the vernal pulse
on May 25 reaches 3,600,000; a midsummer one on August 10,
5,400,000; and the autumnal one appears twice, once on September
21 at 28,800,000, and again on October 19 at 25,200,000. In 1898
the vernal pulse appears May 10 at 1,800,000; midsummer ones, on
July 19 at 10,800,000, and August 9 at 36,000,000 ; and the autumnal
on September 9 at 8,100,000. As in some other organisms, these
pulses are separated by intervals of three to six weeks. The
optimum temperatures lie above 60, though development begins
before that temperature is reached, and the impetus of the autumnal
pulse, or acclimatization to lower temperatures, carries the species
beyond this limit into temperatures of 45. There is a marked
absence of pulses below 60. This seems to be a summer planktont
with no marked preference for the lower temperatures of spring and
autumn.
Scenedesmus quadricauda (Turp.) Breb.* Average number,
9,276,923 (silk, 8,611). In this species, as in the case of others of
the genus and of the Chlorophycecs generally, the numbers present in
1897 were much greater than in 1898 (32,492,647,* silk, 5,818).
Prolonged low water and concentration of sewage afforded stable
32
conditions and food requisite for such development. This species
appears in our collections in every month of the year, though in
much smaller numbers and less frequently from November to
April that is, below 50. Pulses of noticeable magnitude appear
only above this temperature, and usually above 60.
Slight traces of vernal and autumnal pulses appear in the col-
lections of the silk net in 1894-1896. In the filter-paper collections
of 1897-1898 they are well defined. The vernal pulse appears in
1897 on May 25 at 46,800,000, and in 1898 on May 10 at 70,200,000.
The autumnal maximum in 1897 is remarkable both for its large
numbers and its prolongation, culminating twice first on Septem-
ber 21 at 151,200,000, and again on October 19 at 154,800,000.
This remarkable development, combined with the stable conditions
and higher temperatures (Pt. I., PI. XI.) of that low- water autumn,
is responsible for the continuance of the species in our collections
throughout the winter. In 1898 the species declined earlier, in
November, and was but sparingly represented in collections of the
winter of 1898-1899. As in other species of the genus and other
Chlorophycece, midsummer pulses appear at intervals, often of four
weeks, but ranging from three to six. In 1897 these occurred on
July 14 at 55,800,000 and on August 31 at 21,600,000. In 1898
they appear on June 28 at 10,800,000, on July 19 at 79,200,000, on
August 9 at 39,600,000, and on August 30 at 54,000,000. At inter-
vals between the pulses the numbers decrease, and in the regular
collections of 1898 the minima between the pulses do not in any case
exceed 30 percent, of the adjacent maxima, and are usually very
much less. The distribution of the pulses of this species coincides
very closely with that of the other species of the genus, and also with
that of other Chlorophycea. For example, Pediastrum pertusum,
the most abundant of the larger algas, has seven of its thirteen pulses
on the same dates with those of Scenedesmus quadricauda and three
others on adjacent dates, leaving but three which are not practically
coincident. The operation of some common and general factor in
the environment is suggested by such phenomena.
The wide seasonal range of this organism gives it a' claim to rank
as a perennial planktont, though its quantitative distribution
shows clearly that the optimum temperatures for its growth lie
above 60. The largest number recorded in 1897 appears October
19 at a temperature of 65, and in 1898 on July 19 at 84. It is
33
thus predominant only during the warmer part of the year; and
while autumnal and vernal pulses occur, there is no sustained mid-
summer minimum intervening between them. The pulses in
Scenedesmus as a rule follow the volumetric pulses as shown in silk-
net catches (Pt. I., PI. XI. and XII.). Thus in 1897, on September
14 and 21, the plankton measures 19.8 and 3.0 cm. 3 per m. 3 , respec-
tively, Scenedesmus quadricauda numbering 20,700,000 and 151,-
200,000; and, again, on October 5 and 19 the plankton measures
12.92 and 1.86 cm. 3 , and this alga numbers 93,600,000 and 154,-
800,000. Its share in the volumetric pulses is thus indirect to a
large degree, and is perhaps modified by food relations.
Schroederia setigera (Schroder) Lemm.* Average number,
21,450,000. In 1897, 69,040,912. It appears in all months of the
year and in almost every collection. It has well-defined vernal and
autumnal pulses separated by the summer period, in which only
minor pulses occur. In 1898 midwinter numbers are as high as
those of midsummer. Schroederia is thus truly a perennial plank-
tont. The vernal pulse appears in 1897 on April 27 at 302,400,000,
and in 1898 on May 3 at 150,000,000. The autumnal pulse in 1897
culminates on September 21 at 565,200,000, and is followed by sec-
ondary culminations on October 26 at 136,800,000, and on Novem-
ber 23 at 203,400,000. In 1898, when hydrographic conditions
were less stable, the autumnal pulse reached only 50,400,000, on
September 6. This is followed by minor pulses, declining to a mini-
mum in the following February. It disappeared in the collections
with the flood waters of March, 1899. The sequence of these sec-
ondary pulses follows much the same course as has been described
for other species, namely, maxima at intervals of approximately
a month (two to six weeks) separated by more or less sharply
defined minima. There are twelve such pulses (including the major
ones) in 1898 and an interval of seven weeks in March- April in
which none occurs. Six pulses appear in the last five months of
1897.
The optimum temperatures as indicated by the position of the
vernal (60 in both 1897 and 1898, as shown in Table III., Pt. I.)
and autumnal (71 in 1897 and 79 in 1898) pulses lie between 60
and 80. This appearance of the vernal pulse at a lower temperature
than the autumnal (usually about 10 lower) is not confined to this
species but is a general phenomenon among other Chlorophycea.
34
It is apparently a phenomenon of seasonal acclimatization, by virtue
of which the low temperatures of the winter lower the optimum for
the vernal pulse, and the high temperatures of the summer raise
it for the autumnal pulse.
Selenastrum bibraianum Reinsch.* Average number 519,235.
Recorded only from the beginning of August till the end of Novem-
ber, and never in great abundance. Slight evidence of a September
pulse.
Some other Chlorophycecs have been included in the totals as
"unidentified," and isolated occurrences of the following have been
noted: Cerasterias longispina (Perty) Reinsch, C. raphidioides
Reinsch, Dactylococcus infusionum Nag., Glceocystis gigas (Kiitz.)
Lagerh., Staurogenia lauterborni Schmidle, and a few of the Con-
fervacecz which are probably adventitious. These are a species of
Conferva, of Prasiola, and of Ulothrix all of which appear sparingly
in spring and autumn planktons, the first-named and the last
as minute filaments in the filter-paper collections. A thorough
analysis of the unidentified forms would greatly extend the list of
species and varieties.
BACILLARIACE^E.
(Plates I. and II.)
Average number, 396,192,716, including, without duplication,
diatoms from both silk and filter-paper collections. They were
almost twice as abundant in the more stable conditions in which the
collections of 1897 were made. The Bacillariacece are more abun-
dant than any other synthetic group of organisms in our plankton.
They exceed (in 1898) the Schizophycecz five to one, the Chloro-
phycecs seven to one, the desmids eight thousand to one, and the
synthetic Mastigophora by more than four to one. Their numerical
preponderance is, with the exception of the synthetic Mastigophora,
equaled or exceeded by their relative quantitative significance in the
ecology of the plankton.
They appear without exception in every collection, and their
seasonal distribution in its main features is repeated from year to
year. There is a principal vernal pulse in April-May and a hiemal
pulse in November-December. Minimum periods separate these
pulses and are varied by other pulses, usually of minor importance,
at intervals, in 1898, of three to five weeks. The winter minimum
35
is at a lower level than the summer one. In. 1894 the interval of
collection is too great to follow the seasonal distribution, but there
are hints of summer and autumnal pulses. In 1896 there were no
May collections, and the largest number, 6,060,665, appears June
19, five minor pulses on July 18, August 21, September F2, October
11, and November 5 intervening before the hiemal pulse of 3,574,-
028 appears on November 27. Other pulses follow on December 18,
January 6, February 4, March 4, and March 17, before the vernal
pulse of 1896 culminates at 105,440,858 on April 24. This is fol-
lowed by minor pulses on May 18, June ] 1, July 18, August 8, and
September 16, and by the hiemal pulse of December 3 of 346,982,-
928*. The vernal pulse of 1897 appears April 27 at 6,207,473,520,
but is surpassed by a pulse on July 14 principally of Melosira
spinosa of 11,459,289,600, and minor pulses then follow on August
17, September 29, October 26, and December 7 and 21. The hie-
mal pulse of this year is insignificant. In 1898 three minor pulses
appear, January 21, February 15, and March 22, and the vernal
pulse culminates May 10 at 3,865,257,360. Minor pulses follow on
June 14, July 19, August 9, August 30, September 27, October 25,
and November 22, and the hiemal pulse culminates December 15 at
436,535,790, followed in 1899 by minor ones on January 10, Febru-
ary 14, and March 14.
Some of the pulses here indicated are due to the development of
single species, as that of Melosira on July 14, 1897. Most of them,
however, are composite, including a number of species. This is
especially true of the vernal pulse, w T hich in 1898 is due to the com-
bined increase in Fragilaria virescens and F. crotonensis, Cydotella,
Asterionella, Navicula spp., and Synedra acus. Asterionella culmi-
nates early in the vernal pulse and the majority of the others
towards its close. Melosira varians is among these, but M.
spinosa contributes less' to this pulse than it does to later ones.
Minor pulses are also composite, as, for example, that of August 9,
1898, which is due to Melosira spinosa, Cydotella, and Navicula.
FACTORS CONTROLLING DIATOM PRODUCTION.
The fact that many of these pulses represent the combined
fluctuations of a number of species leads us to look for some factor
* Filter-paper collections included in this and in following years.
(4)
36
in the environment common to them all to which these pulses may be
attributed. On the following page the seasonal distribution of the
total diatoms has been plotted for 1898, along with that of the ni-
trates and of the total plankton (volumetric), the thermograph, and
the hydrograph. An examination of the changes in nitrates yields
no marked evidences of correlation. The vernal pulse of diatoms
follows the high nitrates of winter and spring, and the hiemal pulse
in December appears after their autumnal rise, and in this particular
year develops at the- time of an unusual drop in nitrates (Pt. I.,
PI. XLV.). The diatom pulses do not show any constant relation
to the movement in nitrates either in amount or direction. Whipple
('94) has noted the importance of nitrates in the development of
diatoms in reservoir waters. The fact that little correlation
appears in our waters between the fluctuations of the nitrates and
the growth of diatoms may be due to the presence here of nitrates
owing to sewage contamination far in excess of the demands
which the diatoms make, and the limitations placed by other elements
in the environment are reached before that of the nitrate food-
supply becomes operative. The distribution of these diatom pulses
throughout the whole year, even in seasonal extremes, seems to pre-
clude the factor of temperature as the immediate cause of the
pulses except as it may affect the growth of individual species,
which is sometimes apparently the case, as is shown in subsequent
pages.
The vernal pulse is attained each year about May 1, at which
time the water passes the temperature of 60. The average of the
recorded surface temperatures of 1898 in the river is about 58.
Surface temperatures, except in winter months, are usually several
degrees higher than bottom temperatures (Pt. I., Table III.). Our
records are always of diurnal temperatures. The true average tem-
perature, owing to colder water at lower levels and to the nocturnal
decline, will lie several degrees below 58 probably about 55.
The greatest development of diatoms thus takes place at a temper-
ature a few degrees higher than the average temperature for the
year. Owing to the somewhat greater abundance of diatoms dur-
ing the warmer months, the average thermal exposure of the plank-
ton diatoms will be somewhat higher than the average temperature
of the year. There may be some significance in this phenomenon
of the occurrence of the optimum temperature for development at
37
Fig. A. Diagram showing the seasonal distribution of diatoms, total plankton,
nitrates, and thermograph and hydrograph of Illinois River at Havana for 1898.
38
approximately that of the average thermal exposure. The vernal
pulse may, in part at least, be the result of a process of natural
acclimatization. The fact that a similar development does not
recur when this temperature is repassed in the autumnal decline
militates, it is true, against the potency of this temperature as a
factor in the vernal pulse. This temperature is passed in October
(Pt. I., PL VIII. XIII. ), but October pulses are rarely so pronounced
as those of adjacent months. Other factors more potent than tem-
perature are operative at that season of the year.
As will be seen in the diagram, the most pronounced and pro-
longed minimum appears in January, February, and March. In
these months but a single record in excess of 100,000,000 per m. 3
is found. This or at least the first two months of it is the period
of the ice blockade (Pt. I., PI. IX.-XIIL), during which the aeration
of the water by the wind is prevented, and the customary equilib-
rium in gaseous contents may be disturbed. It is the time when
stagnation most threatens disaster to the plankton. The earlier
stages of this blockade in December do not seem to be deleterious
to the growth of diatoms, since at such times the blockade is less
complete, the exclusion of light by the ice less effective, and the
accumulation of the products of decay less pronounced. The data
at hand do not suffice to elucidate the matter further.
The position of the diatom pulses with respect to the movement
of the hydrograph is suggestive though not conclusive of a pos-
sible correlation between the two phenomena. The double vernal
pulse of April-May appears in the declining waters of the major
spring flood. The diatom pulse of June 14 is found in the decline
of the May- June flood. The pulse of August 9 is caught on the ris-
ing waters of a slight flush of the river, and that of August 30 on its
decline. That of September 27 appears after a series of slight rises,
and those of both October and November attend rising water, but
the well-developed pulse of December appears with its decline.
There are, counting the double vernal pulse, ten pulses in 1898,
from March to January. Of these, seven are found on declining
floods, and .but three on rising water, and two of these three appear
during the slow rise of October-November. Furthermore, the
magnitude of the flood is correlated with that of the diatom pulse.
The vernal pulses of 3,453,778,080 and 3,865,257,360 attend the
major spring flood, culminating April 2 at 18 feet; the pulse next in
39
size, that on June 14 of 1,039,619,680, attends the decline of the
flood next in importance that culminating May 25 at 13.9 feet;
while the third pulse, that on December 15 of 436,535,790, attends
the decline of the flood culminating November 25 at 8.7 feet. The
hydrograph of 1897 (Pt. I., PI. XI.) is unlike tha^of 1S98 (Pt. I.,
PI. XII.) in the delay of the so-called "June" rise, which culminates
July 5 at 7.5 feet. Its decline runs through the month into August.
The diatom pulse attending the "June" rise of 1897 appears about
a month later than it did with the earlier pulse of 1898, culminating
July 14 at 1 1,459,289,600. A delay in the flood is thus attended by'
a delay in the diatom pulse. In 1897 there is no December rise and
no diatom pulse of noticeable magnitude, though in 1895, in similar
absence of the flood, there is a well-defined diatom pulse. In 1896
there is a series of five floods, each involving the early stages of
overflow (Pt. I., PI. X.), and on the decline of each occur one or
more diatom pulses.
It is but natural that the greater number of diatom pulses should
fall on declining river-levels, since, as I have previously shown,
these periods exceed in duration those of rising floods. They also
predominate during the prevalence of seemingly favorable temper-
atures, and are characterized by relatively more stable conditions
in the environment. There is, however, it seems to me, another
and more potent reason why diatom pulses appear at such times.
It lies in the overflow of seed-beds in the margins of the permanent
backwaters and the run-ofl of the plankton which develops there
with the fall in levels. This is very apparent to one familiar with
the locality. During the decline of the flood the channel current is
often diverted in minor lateral channels, such, for example, as that
(Pt. I., PI. II.) which courses through Thompson's Lake Slough into
Thompson's Lake and out again into the river at its southern end by
way of "the swale" and the "cut road." A similar current on the
eastern bottoms, which enters partially by way of Mud Lake Slough,
rejoins the river through Quiver Lake. These lateral currents are
joined by the run-off from overflowed bottoms and adjacent
marshes and swamps, all of which, as well as the permanent back-
waters thus draining into the channel, breed at such times an abun-
dant plankton including diatoms. The contributory function of the
backwaters to the plankton of the river proper is thus at its maxi-
mum during the decline of the flood.
As the flood recedes, relict pools on the bottom-lands and along
the margins of the permanent backwaters are formed, in which the
conditions favoring sporulation or other means of providing for
resuscitation are to be found. The emerging bottom-lands thus be-
come the seed-bed for starting a new cycle of diatoms whenever flood
conditions return. In the river, on the other hand, the conditions
for sporulation are not so favorable, and the current tends to carry
away such resting stages as may be formed. The observed facts
regarding the distribution of diatoms and the examination of the
conditions under which these pulses occur thus alike yield corrob-
oration of the view that floods are potent factors in determining the
occurrence of diatoms in fluviatile waters, especially where back waters
are extensive.
The nature of the action of floods is in some respects similar to
that of the overturning of the water which occurs in lakes when the
point of maximum density, 39.2, is passed in either direction. In lakes
of some depth the vertical circulation of so large a volume of water
results in a stirring up of the bottom deposits containing the resting
stages of diatoms, so that they are brought again into increased
light and to better aeration. Whipple ('94) has emphasized the
importance of this overturning in starting the growth of diatoms.
In our shallow waters this physical phenomenon is of less impor-
tance than in the deeper waters of the lake or reservoir. The vol-
ume in circulation is smaller, though some compensation for this
may exist in the possibility of repeated over turnings with fluctua-
tions in temperatures at the critical stage. The existence of cur-
rents, the movements of fish, and the roiling effect of strong and
long-continued winds upon our shallow backwaters, combined with
the fact that much of the seed-bed area of overflow is dry land at
the time of the autumnal overturning, all serve to minimize the
effect of this overturning in our waters upon the growth of diatoms
in the plankton. The spring overturning occurs early in March,
and in 1896, 1898, and 1899 a slight pulse not exceeding an increase of
100 per cent, follows the overturning within an interval of a fortnight.
The vernal pulse is about two months later than the overturning,
and the relation of this to the overturning does not seem to be inti-
mate. The autumnal overturning occurs towards the middle or end
of November, and in 1895, 1896, and 1898 the hiemal pulse of
December follows close upon it, within two, or at most three, weeks.
41
The relation is here more apparent, but the resulting pulse is no
larger than those following upon floods during summer, and but
little larger than the ones which precede it in the autumn. The
effect of this overturning upon the plankton of the Illinois River
may thus be detected, though it is here of less importance than in
lakes and reservoirs since it is overshadowed or replaced by other
and more potent factors.
The relation of the seasonal distribution of the diatoms to that
of the total plankton is not readily unraveled. The latter is the
resultant of a most complex series of factors, whose number and
relative potency are subject to constant change and readjustment
in the unstable environment of the stream. It is the biological
expression of the state of tension among these various factors which
for the moment exists. Of these factors the diatoms are but one,
though an important one, in the food cycle and ecology of the
plankton. The volumetric determinations in. the diagram (p.
37) do not give" the true seasonal distribution of the total plankton
owing to the escape of an unknown quantity through the meshes of
the silk net. They represent more truly that of the animal plank-
ton than that of the phytoplankton. A comparison of the seasonal
distribution of the diatoms and total plankton may serve, in spite
of the errors involved in the volumetric determinations and the
disparity of individuals among the diatoms, to throw some light on
the effect of the fluctuations of the latter upon the movement in the
volume of plankton. A close comparison of the two seasonal curves
reveals the fact that the diatom curve is not identical with the vol-
umetric curve. It is true that the double vernal (April-May) pulse
of diatoms coincides in location with the vernal volumetric pulse.
This is also true of the pulses of June 14 and July 19. The crest of
the volumetric vernal pulse is, however, lodged between the double
apices of the diatom curve, and all the subsequent volumetric
pulses from July on lie in depressions of the diatom curve, and vice
versa. It is apparent at once on examination of our planktons that
the catches of the silk net are from the volumetric standpoint
largely, indeed overwhelmingly, of animal origin. These volu-
metric pulses are as a rule largely pulses of the zooplankton. It is
therefore to be expected that the diatoms would decrease at such
times, since they form the food of many Entomostraca and not a few
Rotifera. The appearance of the diatom pulses before or after the
42
volumetric (animal) pulse may therefore in a measure present the
wavering tendency to establish an equilibrium between these two
elements of the plankton. The presence of an abundant animal
plankton may therefore be a cause of some of the minimum periods
between diatom pulses. Other causes, such as decline of food ele-
ments, may also arise, but in our waters the nitrates at least rarely
ever reach a level where an unutilized margin capable of support-
ing a large diatom population is not still present. Data concerning
other food elements are not at hand, but their paucity in water
derived from such varied sources and so liberally fertilized by
organic wastes seems improbable. There is also the further possi-
bility and, indeed, from the data in hand the probability of the
existence among diatoms of reproductive cycles, interrupted by
resting periods. The available data do not, however, throw any
light upon the nature of this internal factor or the cause for the
running down of the energy of reproduction, and but little upon the
operation of environmental factors which stimulate anew the
process of reproduction.
The seasonal distribution of the diatoms as a whole, and that of
individual species also, offer repeated instances of recurrent pulses
at intervals approximating four weeks the lunar month. In 1898
thirteen such pulses can be detected. These often correspond
roughly to minor flood intervals, but not always so, for occasionally
two pulses occur on the decline of a single flood. Similar appear-
ances may be traced in other years, when collections were frequent
enough to exhibit minor pulses. They are, however, in all cases
quite irregular, and exceptions are frequent.
That cosmic factors may indirectly, through immediately environ-
ing factors, affect the reproductive phenomena of pelagic organisms
has been suggested by the work of Kramer ('97), Mayer ('00), and
Friedlander ('01) in the case of the "Palolo" worm, a coral-reef
annelid whose seasonal swarming for reproductive purposes occurs
at somewhat definite lunar intervals.
While the data concerning the seasonal distribution of diatoms
in the Illinois River may serve to suggest the operation of an enig-
matic cosmic factor, I wish distinctly to state that in my opinion
they are wholly inadequate to establish either its presence or its
potency. It is much more probable that we have to deal merely
with some matter of food relations between the plants and animals
43
of the plankton, and perhaps with the result of increased photo-
synthesis in periods of lunar illumination, which tends to establish the
limits of the pulses.
The number of forms of diatoms noted in our records in the
plankton of the Illinois River is thirty-one. This number could be
greatly increased by the inclusion of the many adventitious species
which flood-waters bring into the plankton and by the addition of
rarer limnetic species. Of these thirty-one at least twelve are
eulimnetic, while the others are in the main adventitious. There
are no species among them peculiar to the potamoplankton, and
the dominant forms here are also abundant in the fresh-water plank-
ton of our own Great Lakes and of European streams and lakes,
barring a few mooted points of specific identity.
The limnetic species are fourteen in number, viz. : Asterionella
formosa, A. gracillima, Cydotella kuetzingiana, Diatoma elongatum
var. tenue, Fragilaria crotonensis, F. virescens, Melosira . granulata
var. spinosa, M. variant, Meridian circulare, Rhizosolenia eriensis,
Stephanodiscus magaroz, Synedra acus, S. acus var. delicatissima,
and Tabellaria fene strata. Of these limnetic forms the more impor-
tant ones are Asterionella gracillima, Cydotella, Fragilaria virescens,
Melosira granulata var. spinosa, and Synedra acus and its varieties.
The absence or small number of certain limnetic species is notice-
able. These are several species of Tabellaria and Attheya. On ac-
count of the abundance of silt and the transparency of Attheya it
may have been overlooked. It has hitherto been reported from
waters much nearer the sea, and this coupled with its affinities to
marine diatoms may explain its absence in our waters.
The remainder of the forms are adventitious, or largely so, and
with the exception of the species of Navicula they have little effect
upon the ecology or quantity of the potamoplankton.
DISCUSSION OF SPECIES OF BACILLARIACE^I.
Asterionella formosa Hassall. Average number of individual
cells, 960. Average size of colony, 4.8 cells. Recorded only in
November, December, and from February through April, and never
in large numbers. The greatest pulse attained at any time cul-
minated on March 30, 1896, at 54,540. Aside from an isolated
occurrence on June 27, 1896, no individuals were recorded at tem-
peratures above 48, and three fourths of the occurrences are at
44
temperatures below 40. The data .are insufficient to trace the
pulses satisfactorily. This species is distinguished with difficulty
from A. gracillima, and may include only old, and in our planktons
often heavily incrusted, individuals; or it may be only a low-tem-
perature variety of the species above named, which in the grand
total of all our collections outnumbers it ten thousand to one.
Asterionella gracillima Heib. Average . number of individual
cells, 28,860,160. In 1897 the species was only one third as abun-
dant, a contrast which finds its explanation in the fact that the
June rise of that year (Pt. I., PI. XI.) did not reach the stage of
overflow, and a June pulse is absent in the collections of that year.
The seasonal distribution of this organism is one of the best-defined
and most striking of all the components of the river plankton. It is
peculiar in the fact that it appears in numbers only during spring and
the beginning of summer, and in the absence of any autumnal pulse
upon the return of the temperatures in which the spring pulse ap-
peared. This species was recorded in every month of the year but
October, but always in small numbers after July 1. In 1894, collec-
tions were not commenced until after the time of the spring pulse.
In 1895 the spring collections were few, and at intervals so great as to
preclude the detection of the full course of the spring pulse. The
maximum number in the collections 'of that year appears April 9 at
1,203,100 and falls to 445,995 on April 29 which is approximately
the time of the maximum of subsequent years. This was a year of
unusually low water during the spring, and overflow stage was at no
time reached (Pt. I., PL IX.), which may account for the apparent
suppression of the spring pulse. The species does not reappear in the
collections of that year until December, but it continues in small
numbers (less than 5,000 per m. 3 ) until the end of March, 1896, when
there is a rapid increase which culminates April 24 at 26,281,400. It
disappears entirely from the records at the end of a fortnight, and save
for a single entry in June and two in September it does not again
appearin 1896. In 1897 the culmination of thespring pulse occurs April
27 at 324, 633, 600 three hundred-fold larger than in the previous
year. There is a normal March flood (Pt. I., PI. XL), on the declining
stages of which this pulse appears. With the close of June the
species disappears from the records. The June rise does not reach
the stage of overflow, and the scanty records show but this single
pulse throughout the year. Beyond a single entry in August and in
45
November the species does not again appear in the records during
the year. In 1898 there is an unusual midwinter pulse on January
11 of 146,280, followed by a decline and irregularities due to the ris-
ing winter flood (Pt. I., PI. XII.). At the middle of March a rapid
increase ensues, culminating April 26 at 891, 648,000 ^on-the declin-
ing spring flood. A decline to 197,683,200 is found at the close of a
week, and it is accelerated by the secondary spring flood, which
attains the overflow stage of 15 feet in the closing days of May
(Pt. I., PI. XII.). With the decline of this flood in June a second
pulse appears, increasing from 15,080 on May 26 to 336,194,880 on
June 14, and at the end of three weeks the species practically dis-
appears from the plankton. A few scattered entries appear during
the summer and fall, and a minor pulse of 10,500 appears on Decem-
ber 20, followed by a decline in the next month.
This species in our waters exhibits a well-defined vernal pulse
towards the end of April at about 60, but no autumnal pulse
appears when this temperature recurs. , There is a slight indica-
tion of a minor midwinter pulse at the minimum temperatures
of the year. This occurrence of a midwinter pulse was noted
by Whipple and Jackson ('99) in the reservoirs of the Brooklyn
water- works, and in the same paper its seasonal distribution in
Fresh Pond, Lake Cochituate, and Wenham Lake, Massachusetts,
is given for the years 1890-97, in the majority of which a mid-
winter pulse commensurate in magnitude with the vernal pulse is to
be found. Autumnal pulses are of infrequent occurrence, the vernal
pulse being the most frequent but not constant. In European
waters no such long-continued examination of the seasonal distribu-
tion of this organism has as yet been reported. Apstein ('96) finds
two pulses per year in Ploner See in May and the last of July ; and
two in Dobersdorfer See, one in April and one in October, separated
by midsummer and midwinter minima. Lauterborn ('93) finds that
this species in the " Altwasser" of the Rhine attains its maximum in
June and again increases in October. In the backwaters of the Elbe,
Schorler ('00) reports Astenonella as abundant in April, June, July,
and October, but refers the organisms to the preceding species. The
existence of the vernal pulse only in our waters is thus somewhat
unique, and the cause of the phenomenon probably lies in some
environmental conditions, perhaps in our peculiar bacterial and
sewage contamination of the autumn. Our vernal pulses appear on
46
declining floods about the end of April at about 60. It can not be
temperature which limits the occurrence of the species, for this
apparent optimum recurs again in October. This is the period of
declining nitrates (Pt. I., PL XLIII.-XLV.), but they rise again in
the autumn, and in our sewage-fed waters they contain even in the
midsummer minimum a quantity adequate to support an abundant
growth of Asterionella. Whipple and Jackson ('99) have found on
analysis that Asterionella to the number of 10,000,000,000 per cubic
meter yield but .079 parts per million of organic nitrogen. The nitrates
in our waters rarely fall below. 25 parts permillion, which, with the other
forms of nitrogen that may be available, would seem to afford nurture
not only for Asterionella but also for competing organisms. These
authors have also found that silica to the amount of 1 . 78 and manganic
oxide to .03 per million are contained in Asterionella to the number
per cubic meter above quoted. As was shown in Pt. I., p. 234, the
silica is present in great excess (26 to 81 parts), and the manganic
oxide, though not reported in the analyses of November waters, is
present on June 15 to the amount of .07 parts per million more than
double the amount required to support Astenonella to a maximum
twelve times as great as any recorded in our plankton collections.
This also occurs at a season when Asterionella is usually declining
rapidly in numbers. Such chemical data as are available thus afford
us no explanation of the limitation of Asterionella in our waters to
the vernal pulse alone.
Some evidence bearing on a factor which may be operative in
producing this phenomenon is to be found in the hydrographic con-
ditions attending the vernal pulse. As previously noted, this
appears, each year with the decline of the spring flood. A repetition
of the overflow in 1898 at the end of May brought with it a repetition
of the vernal pulse of Asterionella in early June. With the decline
of the flood the backwaters make their major contribution to the
channel plankton, and it is during this period that Asterionella
reaches its maximum and also declines. If the spring flood is sup-
pressed, as in 1895 and 1896, the spring pulse of Asterionella is cor-
respondingly feeble. The environmental conditions are thus more
favorable in the impounded backwaters than in the main stream.
Whipple and Jackson ('99) have noted in frustules of this diatom
the appearance of structures which they interpret as spores. If these
are spores, and if the sedimentation of spore-bearing frustules occurs
47
extensively in the relict pools of the emerging bottom-lands, a seed-
bed for re-stocking the waters of overflow is formed with each declin-
ing flood, and this seed-bed becomes potent only when floods return.
The absence of an autumnal overflow and the minor part that the
autumnal overturning plays in our shallow waters whm 39.2 is
passed, may alike tend to suppress here the autumnal or midwinter
pulses which occur elsewhere in deeper water.
The occurrence of the vernal pulse of Asterionella in the last days
of April brings it into close relation with the major volumetric pulse
of the year (Pt. I., PL IX.-XIL). It is not only an important con-
stituent of this spring maximum, but it is one of the most prominent
primal sources of food of the Entomostraca Bosmina, Daphnia,
Cyclops, and Diaptomus, all of which exhibit an increase in numbers
at this period. It shares with Cyclotella the claim to the first place
quantitatively among the synthetic organisms upon which the
early spring plankton depends for its development.
Our records are all based upon the catches of the silk net, through
whose meshes the isolated cells of Asterionella readily escape. Filter-
paper catches give much higher numbers except during the period of
maximum, when the numbers by the two methods do not materially
differ. This seems to be d.ue to the fact that isolated cells are rela-
tively much more abundant after the maxima than they are be-
fore them, and especially at the time of their appearance. These
diatoms form arcs, circles, or whorls, of a varying number of cells.
During the vernal pulses of 1898 the average number in these clus-
ters in the middle of March was three or four, and at the time of the
maximum on April 26 it rose to five or six, often reaching sixteen or
more. A fortnight after this maximum the average fell to 1.4, rising
again with the second pulse, on June 14, to 8.4, and declining in three
weeks, with the fading out of the pulse, to 1.2.
Asterionella is frequently infested with great numbers of a minute
craspemonad flagellate protozoan which appears in thick-set rows
upon the ray-like cells, a single cell sometimes bearing a score of
these organisms. This diatom exhibits considerable variation in size
and proportions. The longer and more slender cells appear at the
times of the maxima.
Cocconeis communis Heib.* Average number, 520,000, but more
than three times as abundant in 1897. This diatom occurs some-
what irregularly in the filter-paper collections, and has been recorded
48
in every month of the year. It is somewhat more prevalent in
spring and autumn, and there are indications of a vernal pulse in
May and an autumnal one in September, separated by prolonged
midsummer and midwinter minima. Vernal pulses appear in 1897
on June 28 at 14,400,000, and in 1898 on May 17 at 7,200,000. Autum-
nal pulses occur in 1896 on September 16 at 2,700,000; in 1897 on
September 29 at 10,800,000; and in 1898 on September 13 at
5,400,000. The optimum temperatures lie between 60 and 75, the
autumnal pulse appearing in higher temperatures than the vernal as
a rule. This diatom is reported as often epiphytic upon algae, and
it may be wholly adventitious in the plankton. There is nothing,
however, in the curve of its distribution to corroborate this view.
Cydotella kuetzingiana Thw.* Average number 243,659,615,
but slightly more abundant in the preceding year. This is one of the
smallest as well as one of the most abundant of all the diatoms of the
river plankton. It readily escapes through the meshes of the silk
net, and plankton collections made by this means give no adequate
conception of its prevalence or importance in the ecology of the
plankton. It appears in every month in the year and in practically
all of our collections, and is thus a perennial planktont. There is a
considerable variation in size among the individuals in the plankton,
but the greater number lie near the smaller rather than the larger
limits. It may be that several species have been combined in the
enumeration.
The fluctuations in the seasonal distribution of this diatom are
considerable, and pulses occur at all seasons of the year. The vernal
pulse is, however, preeminent, and is not approached in magnitude
by those of any other season of the year. In 1897 this pulse culmi-
nates at 5,724,000,000 on April 27, and in 1898 on April 26 at 2,880,-
000,000. Throughout the summer and autumn in both years there
is a series of minor pulses at intervals of two to eight weeks. In 1897
an autumnal pulse of 223,200,000 appears on September 29, and
though not of greater magnitude than two previous summer pulses,
it does surpass anything prior to the pulse of the following spring.
In 1898 there are seven pulses during the summer and fall, culmi-
nating as follows: on May 10 at 2, 668, 000,000; on June 28 at 291,-
000,000; on July 19 at 561,600,000; on August 9 at 401,400,000; on
August 23 at 122,400,000 ; on September 6 at 1 15,200,000 ; on Septem-
49
ber 27 at 57,600,000; on October 25 at 25,200,000; and in December
a pulse well sustained throughout the month culminates on the 15th
at 414,000,000.
The temperature optimum appears to be about 60, though its
return in the autumn does not induce a development comparable
with that of the closing days of April. The midsummer pulses and
that of December show that other causes than temperature are
operative in regulating the occurrence of this organism.
The appearance of the vernal pulse of Cyclotella at the time of
the volumetric maximum (Pt. I., PL IX.-XII.) in April-May sug-
gests its function as one of the primal sources of food for the animal
components of that plankton. The plates are based on collections
of the silk net, and Cyclotella constitutes an insignificant part of the
volumetric total there graphically presented, since it is so small that
it escapes readily through the silk.
Cymatopleura solea (Breb.) W. Sm.* Average number, 2,115
(silk, 1,292), but slightly more abundant in 1897. Isolated occur-
rences in small numbers appear during the colder months, generally
below 60, though several individuals appear in summer records.
This is apparently an adventitious planktont, whose presence is
often due to flood waters.
Diatoma elongation var. tenue Van Heurck.* Average number,
2,471,923. This is a perennial limnetic diatom occurring in every
month of the year and in the majority of our collections. It is but
sparingly present during midsummer. There are well-defined
vernal pulses in 1897 on May 25 of 50,400,000, and in 1898 on May
3 of 18,000,000. A second large pulse appears on the approach of
winter, in 1897, on November 15, culminating at 2,700,000, and in
1898, on November 22, at 9,000,000. In the silk collections of 1895
and 1896 pulses also appear in the last days of April and in Novem-
ber or December. The records thus indicate a decided preference
of the species for temperatures below 70 and the possibility of
rapid development in midwinter as in 1895, during a fortnight of
minimum temperatures (32 + ), culminating at 53,424 (silk) Decem-
ber 18. The vernal pulses coincide approximately with the volu-
metric maximum, and the December pulse of 1895 attends an
unusual winter development of the plankton (Pt. I., PI. IX. and
Table III.).
50
Diatoma vulgare Bory occurred sparingly at irregular intervals,
and is apparently an adventitious species in the plankton.
Encyonema prostratum (Berk.) Ralfs appears a few times during
the summer months, and is evidently adventitious, as is also the
still rarer Epithemia turgida Kiitz.
Fragilaria crotonensis (Edw.) Kitton. Average number of cells,
2.1. This limnetic diatom is much less abundant in our waters
than the following species. In 1898 it appeared in February, and
increased from 19,200 on April 19, to 14,469,120 on May 10, dis-
appearing entirely from the records after May 17. Such meteoric
pulses were not detected in previous years, when only scattered
entries in April, May, and December were recorded. The number
of cells in the filaments is very much less than in F. virescens, aver-
aging but 14 to its 108. Its optimum temperature lies about 60,
and its vernal pulse occurs immediately after the volumetric maxi-
mum (Pt. I., PI. XII.) and upon the same date with that of F. vi-
rescens. It seems to be predominantly a vernal planktont in our
waters. In German lakes Apstein ('96) finds maxima as late as
June-July, but always, it seems, at temperatures below 70.
Fragilaria virescens Ralfs. Average number, 73.1. Apparently
ten times more abundant than in 1897, as a result possibly of the
absence of collections during the period of the vernal maximum in
that year. This is a perennial organism, with two well-defined
pulses ; a vernal one in April-May and another in November-
December. The uniformity with which these pulses appeared in
1895-1898 is very striking when one considers the unstable environ-
ment in which the pulses occur. In 1894 the species is not present
in numbers in any of the scattered collections of the year. In 1895
the vernal pulse is indicated in the collection of April 29 (2,754,675),
after which the species disappears until September, increasing
with a temporary backset by the December flood (Pt. I., PI. IX.)
to a second culmination December 30 at 282, 225. After a minimum
in January, 1896, the numbers increase, with minor fluctuations, to
a vernal maximum of 76,224,000 on April 24, followed by a mini-
mum period from May 18 to the following November. The winter
pulse again appears in December, culminating on the 3d at 867,048.
In 1897 the vernal pulse seems to culminate somewhat later than
usual, though the interval of collection is too great to follow its full
course. The maximum appears on May 25 at 3,549,600, after
51
which the species dwindles away and disappears in August to return
early in November. The winter pulse culminates December 14 at
8,159,250, at a break in the ice blockade. In 1898 the winter mini-
mum continues into April, and the vernal pulse appears May 10 at
253,960,000, rising with rocket-like suddenness from 390,000 of the
previous week, and declining the week following to 4,110,400. The
decline to the summer minimum is prolonged into July, and the
species does not reappear until October. The winter pulse begins
earlier than usual, on November 1, and is well sustained through
the month, culminating on the 29th at 2,254,000. The winter mini-
mum which follows, does not reach the low levels of that of summer.
This species has thus a characteristic distribution, the analysis
of which is by no means simple. The contrast between the summer
and winter minimum may be due to the low nitrates of the summer
and the larger amount in the winter (Pt. I., PI. XLIII.-XLV.),
which favor a proportionate development of this diatom, though
not every species shows this response. The two minima separate
the seasonal occurrences of this species into two periods of growth ;
a vernal, from March to June, and a hiemal, from October to Janu-
ary, the limits and relative development of each being somewhat
variable from year to year. The temperatures of the two periods
differ. Both are times of rapid change, of rise and fall respectively,
and the culminations of the periods of growth lie at widely sepa-
rated temperatures. The vernal pulses in 1896 and 1898 in which
years collections were frequent enough to locate them with some
degree of accuracy appear at 72 (April. 24) and 61 (May 10)
respectively, and in every year the vernal pulse appears during a
period of rapid change. The hiemal pulse, on the other hand, cul-
minates in each year after the winter minimum approaching 32 has
been reached, and in two years during the ice blockade. Tempera-
ture within these limits seems not to be a determining factor in the
pulses of this organism. The nitrates (Pt. I., PI. XLIII.-XLV.)
have been uniformly high (above 2 parts per million) whenever the
pulses occurred. In 1898 they decline abruptly (Pt. I., PI. XLV)
and remain at a low level throughout December, and in this month,
when usually Fragilaria attains its hiemal maximum, we find it
dropping to the unusual minimum of 20,000. The pulse which
began in November is cut off apparently by this unusual decline in
nitrates. Abundance in nitrates is not, however, in itself sufficient
(5)
52
to cause a pulse of development of Fragilana, for nitrates are
abundant when the diatom declines and is at its minimum. It
does not seem possible to find in the unstable environment of this
organism any external factor which shows a causal connection with
its periods of growth.
Apstein ('96) found that this diatom reached its major pulse
in March and April in Dobersdorfer See, and a minor one in
November.
The cells of this diatom form long twisted bands, visible to the
unaided eye. They reach a much greater length in this species
than in the preceding one, and are longest during the height of the
growing period, decreasing rapidly in length as it declines. The
average number of cells in a ribbon at the time of the maximum lies
between 150 and 200, and at other times is usually below 100 and
often below 2 5 .
The vernal pulse of this species coincides with that of F. croto-
nensis, and appears either with or just after the volumetric pulse.
The December pulses may in part serve as primal food sources for
the fairly constant minor volumetric pulse of December.
Gomphonema constrictuvn Ehrbg.* Average number, 501,923.
This species appears irregularly, with a predominance of occur-
rences in May and November, and is apparently adventitious.
Melosira granulata (Ehrbg.) Ralfs var. spinosa Schroder.
Average number of cells, 1,181,125 (filter-paper, 34,762,365).
In 1897 it was more than five times as abundant. In the
filter-paper collections as a whole it is about fifty times as abun-
dant as in those of the. silk net. A much greater proportion
of single cells and short filaments occurs in the latter collections,
since the longer filaments are the more readily retained by the silk.
In the discussion which follows, the data from the silk collections
will be used, since they cover the whole period. The data from
the filter-paper collections indicate very nearly the same seasonal
routine, and the differences between the results by the two methods
lie in the proportions of the numbers rather than in the direction of
movement in the fluctuations. The pictures of the seasonal
changes in occurrence of the diatom given by the two methods are
essentially alike aside from greater irregularity during minimum
periods, resulting from the larger margin of error in the filter-paper
method as I used it.
53
This Melosira is a perennial planktont in that it occurs in every
month of the year in the river. Its appearances from December
to March are, however, irregular, and its numbers small. Its large
pulses above 1,000,000 all lie between May 15 and October 1,
with the single exception of the pulse of April 24, 1896, culminating
at 2,056,400, in temperatures of 72, occurring fully a fortnight
earlier than usual. The major pulse seems normally to occur in
June; at least in 1896 and 1898, when collections were frequent at
this season of the year, such pulses appear on the llth at 12,940,000
and on the 21st at 32,114,880. A June pulse also appears in 1895.
September pulses appear in 1895, on the 12th, at 2,254,182, and in
1898, on the 27th, at 5,499,840. There is, however, no well-defined
vernal and autumnal growth period, since large pulses occur through-
out the whole summer. The greatest pulse on record (111,456,000)
is on July 21, 1897, and in 1898 there are three minor pulses between
those of June and September. Including the major pulses, there
are in 1895 five, in 1896 six, in 1897 five, and in 1898 eight, pulses
at intervals of two to six weeks between May and October, the ones
at either end of the season being often but slightly developed, the
remainder usually running from 1,000,000 to 5,000,000.
This species is predominantly a summer planktont, and its
optimum temperature lies above 70, the greatest number recorded
appearing at 81. This is one of the most abundant diatoms of
the potamoplankton, and in our waters it attains its greatest de-
velopment during the season of the minimum occurrence of nitrates,
in whose utilization it is quantitatively an important agent. It
fills the gap between the vernal and autumnal or hiemal appear-
ances of Asterionella and Fragilaria, thus providing a continuous
source of food for the zooplankton with w r hich it is associated. It
is, by virtue of its numbers, its size, and its seasonal distribution,
quantitatively and ecologically the most important of all the
diatoms of the plankton of the Illinois River.
The only factor in the environment to which the limitation of the
rapid growth of this species 'to the May-October period can be re-
ferred is temperature. There are but three instances in the records
of Melosira exceeding 100,000 per m. 3 at temperatures below 60, and
one of these is but a few days prior to the attainment of that temper-
ature. It cannot be food which deters its development below this
point, since the nitrates at least are then most abundant (Pt. I., PL
54
XLIII.-L.). Other diatoms, as in the hiemal pulse of Fragilaria,
develop in numbers at temperatures approaching 32, but not M.
granulata var. spinosa. Whipple ('94) concludes from the records of
examinations of potable waters in Massachusetts that temperature
has possibly a slight influence on the growth of diatoms, but that it is
of so little importance that it does not affect their seasonal distribu-
tion ; and, on the other hand, that a sufficient supply of nitrates is one
of the most important conditions for their growth. The seasonal dis-
tribution of Melosira was not separately discussed in his paper
though included in his general statements. In our waters the data
at hand seem to show conclusively that abundance of nitrates is of no
avail in the case of Melosira when the temperature falls below 60.
There are times, therefore, in the case of this, our most important
diatom, when temperature is more potent than food as a factor con-
trolling its growth.
Melosira does not appear in its maximum pulses at the time of the
major volumetric pulse of the total plankton of April-May, nor do
its fluctuations seem to bring about directly any considerable
changes in the volume of the plankton. For example, the extreme
pulse of 111 ,456,000 on July 21, 1897, occurs at the time of a sudden
drop in the amount of plankton (Pt. L, PL XL). The amount of
plankton on July 14, 21, and 30 is 8.16, 0.92, and 1.05 cm. 3 per m. 3 ,
and the corresponding numbers of Melosira are 66,528,000, 111,456,-
000, and 13,176,000.
The diatoms here discussed are predominantly of the type
designated as var. spinosa, marked by the spinous prolongations
from the valves at the ends of the filaments. The cells of the forms
in our plankton are proportionately much longer, as a rule, than
those figured by Schroder ('97), usually attaining one and a half to
two times the length without proportional increase in diameter.
Not infrequently in the height of the growing season much elongated
and curved cells and filaments are to be found. In one instance an
unusual number of filaments approaching M. varians in form though
still, of the spinous type were found. It is not improbable that
several so-called species of Melosira have been included with this
variable species in the enumeration.
Melosira is the bearer of numerous passive planktonts, the most
abundant of which is Bicosceca lacustris Clk. Associated with this,
and often on the same filament, is the elegant little craspemonad
55
Salpingceca brunnea Stokes. Cells to which several of these flagel-
lates are attached very frequently exhibit a breaking up of the cell
contents into eight brownish masses, often of spore-like form, and it
is not an uncommon thing to find such parasitized filaments with
several empty cells. The eggs of the rotifer Diurella ligris are fre-
quently found attached to the filaments of this diatom. The num-
ber of cells in the filaments in the silk collections averages 6.4 in 1897,
and 7 in 1898, while in the filter-paper collections it averages 3.5 in
both years. The numbers per filament range from 1 to 40, and the
filaments are wont to be somewhat longer during rapid growth than
in periods of decline or minimum.
Melosira varians Ag. Average number, 148,626 (filter-paper
3,455,538). The discussion is based upon silk catches. The species
was about equally abundant in 1897 but much less so in previous
years. This is a perennial species, reported in every month of the
year and in most of the collections. It exhibits two well-defined
pulses, a vernal one in April-May and an autumnal one in September-
October. The reduction in the minimum intervals varies from sea-
son to season and from year to year. It was most pronounced, al-
most to suppression, in July and August in 1894, 1895, and 1896, and
in December-February in 1896-97 and 1898-99. In other seasons
the minimum falls to 1,000 to 15,000.
The vernal pulse (146,916) appears in 1895 on April 29, in 1896
(229,235) on May 18, in 1897 (2,419,200) on May 25, and in 1898
(3,164,160) on May 5. The autumnal pulse (150,720) is found in 1895
on October 30; in 1896, on September 16 at 378,900; in 1897 there
are two pulses, one on August 30 at 738,000, and the other on No-
vember 15 at 458,800; and in 1898 one, on October 18 at 348,000.
The autumnal pulses are thus much smaller than the vernal ones
and exhibit a greater range in the time of their appearance.
As in the case of many other organisms this diatom also exhibits
the phenomenon of recurrent minor pulses at intervals of a few
weeks. They range in height from 25,000 to almost 1,000,000, and
are largest when found in the proximity of the major pulses. The
records are not frequent enough to trace them in all seasons. They
appear in January in 1896, 1898, and 1899; in February in 1898;
twice in March in 1896; in April in 1896; twice in June in 1897 and
again in 1 898 ; in July in 1897 and 1898 ; in August in 1897 and 1898 ;
56
in September in 1898; in November in 1896, 1897, and 1898; and in
December in 1894.
The optimum temperatures, omitting the pulse of August 30,
1897, at 80, all lie below 72, averaging 65 for the vernal pulse
and 62 for the autumnal. But three pulses in all, exceeding
100,000, lie at temperatures above 70, and but three below 50.
In the case of this species likewise temperatures seem to be potent
factors in limiting its seasonal occurrence. The fluctuations in
nitrates do not seem to bear any constant relation to its develop-
ment. The midsummer minimum of the diatom may appear, as
in 1896, during an abundance of nitrates (0.5 to 3.0 parts per mil-
lion Pt. I., PI. XLIII.) unusual for the season. On the other hand,
a minimum of nitrates (.1 to .35) in August and December, 1898,
coincides with a suppression of this species in the plankton. Thus
in the presence of food, temperature seems to be a determining
factor in the seasonal distribution of this organism. Whipple ('94)
expresses the opinion that the growth of diatoms occurs at those
seasons of the year when the water is in vertical circulation ; that is,
when it passes 39.2. In our waters this generally occurs early in
March and late in November. In this species the only pulses
which it seems might exhibit the effect of this phenomenon are
those of December and March, and neither of them are in any way
constant or prominent. Neither of the major pulses, vernal nor
autumnal, can be attributed to it. The latter pulse occurs prior
to the autumnal overturning of the water.
The vernal pulse usually follows the spring volumetric maxi-
mum, and the autumnal one generally appears during a volumetric
minimum. No immediate quantitative effect of this species upon
the plankton is apparent.
In European waters this is a common planktont, and Apstein
('96) reports vernal maxima in March, April, and May, and an
autumnal one of minor value in November.
The number of cells in the filaments varies from one to sixty,
and in filter-paper collections averages four, while in the silk
catches it varies from seven to fifteen from year to year. The fila-
ments average somewhat longer during the periods of maximum
growth, reaching twelve to twenty-five. This species also occasion-
ally bears the flagellates found upon M. granulata var. spinosa, but
not in such abundance. It is quantitatively much less important
57
in our plankton than that species, though this does not seem to be
the case in some European waters.
Meridian circulare Ag. has appeared but four times in winter
planktons, from December to March, and seems to_ be adventi-
tious.
Navicula iridis Ehrbg.* Average number, 297,307. Appears
at irregular intervals, often with flood waters and in the colder
months. It seems to be adventitious.
Navicula spp.* Average number, 8,569,038. About twice as
abundant in 1897. Under this head I have included a number of
species of Navicula, and, possibly, even species of genera resembling
Navicula. The individuals are all of small size, and are principally of
the type of the smaller forms of N. brebissonii Kiitz. and N. gracilis
Ehrbg. They are quite abundant in collections from Quiver Creek
and Spoon River. Their greater abundance in 1898 as compared with
1897 may be caused by the greater movement in river levels in the
former year (85.6 ft.) as compared with that of the latter (55.5 ft.).
This feature of the distribution of these forms suggests that they
are adventitious in the plankton. This view is further supported
by the fact that some, though not all, of their apparent pulses
appear with flood waters; for example, the pulse of 64,000,000 on
May 17, 1898. There are indications, independent of floods, of
pulses in April-May and November-December, which may, how-
ever, be simply reflections of pulses in the normal habitat of these
diatoms the shores and bottom of the river and its tributaries.
They are represented in the plankton at all seasons, and the diver-
gence in numbers is at no time so marked as it is in typical plank-
ton diatoms, such as Asterionella.
Nitzschia amphioxys (Ehrbg.) Kutz. appeared several times in
winter collections, and N. sigmoidea (Nitzsch) W. Sm. is adventi-
tious in small numbers in flood waters. Several species of Pleu-
rosigma appear at irregular intervals throughout the year in both
flood waters and stable conditions and are apparently adventitious,
appearing in relatively small numbers.
Rhizosolenia eriensis H. L. Smith was noted on a few occasions
in winter planktons. Its exceeding transparency and the abun-
dance of silt and debris at the times of its occurrence so obscure it
that it may have escaped detection in many instances.
58
Stephanodiscus niagarce Ehrbg., a common planktont in the
waters of the Great Lakes, appeared but once, in May, in our plank-
ton, though the river had for years received, by way of the Chicago
River, constant access of water from Lake Michigan. The turbid,
sewage-laden, and warmer waters of the Illinois are evidently not
favorable for its growth.
Surirella ovalis Kiitz. var. minuta (Breb.) Kirchner.* Average
number, 761,538. Present sparingly throughout the year, but
principally during summer months. Vernal pulse in May.
Surirella spiralis Kiitz. Average number, 1,612. Less abundant
in the more stable conditions of 1897. This species is most abun-
dant in Quiver Creek and Spoon River. Its fluctuations are slight,
irregular, and often appear with flood waters, all of which phenom-
ena indicate its adventitious character in the river plankton.
Synedra acus Kiitz.* Average number, 36,558,462 (silk, 308,-
330). This species is a perennial planktont, appearing, for example,
in 1898 in every collection. It has a highly developed and shifting
vernal pulse, and an inconstant and but slightly developed autumnal
or hiemal pulse. The vernal pulse appears in 1895 on April 9 at
209,880; in 1896 on April 24 at 366,828; in 1897 on May 25 at
2,620,800 (82,800,000*); and in 1898 on May 10 at 9,043,200
(813,600,000*). The second pulse appears in 1895 on November
14 at 99,360; in 1896 on December 3 at 44,464; in 1897 no pulse
occurs; in 1898 it occurs on November 8 at 19,000. As in some
other diatoms, there are minor pulses throughout the year, though
in this case they are all feebly developed, exceeding 100,000 (silk)
in but a single instance. The minor pulses of midwinter often
exceed in prominence those of midsummer. The meteoric char-
acter of the vernal pulse is very pronounced in this species both in
the suddenness of its appearance and its disappearance and in the
height which it attains.
The variety delicatissima W. Sm. is included here with the type
acus. During the autumn of 1898 a separate record was kept of
the two, with the result that the variety appears to include about
four fifths of the individuals at that season. The tw T o are not
readily separated. The colorless form recently described by Pro-
wazek ('00) as 5. hyalina is also included, and it is not uncommon
when S. acus is abundant. Colorless forms of other diatoms of the
plankton, as Asterionella, Melosira, and Fragilaria, also occur, but
59
it would seem from the intergradation with the normal condition
that it is a phenomenon of physiological import rather than of
specific significance. It would seem desirable that experimental
breeding of diatoms should be employed as a test before specific
diagnoses utilize this character.
Synedra capitata Ehrbg. is occasionally adventitious in the plank-
ton in spring months.
Synedra ulna (Nitzsch) Ehrbg.* Average number, 302,308
(silk, 34,510). This appears somewhat irregularly in the plankton,
with a vernal pulse on May 17 of 5,400,000 and an autumnal one
November 15 of 1,800,000. It is abundant on the ooze of exposed
springy shores after rapid decline of the river, and is probably
adventitious in the plankton to some extent from this region.
Tabellaria fene strata Kiitz., which is exceedingly abundant in the
plankton of European lakes and in our own Great Lakes, was found
but a single time in the w r aters of the Illinois. It can hardly be lack
of food elements which prevents its development, and there are
times when favorable thermal conditions would seem to be offered
in spring and autumn, when the river temperatures do not exceed
the summer temperatures of our Great Lakes. It may be that the
chemical conditions attending sewage contamination exert a dele-
terious influence upon this species and others of the genus, such as
T. flocculosa, which abound in purer lake waters.
CONJUGATE.
This group of algae is represented in the plankton only by a few
desmids, which neither in number, or quantity, play any important
part in the ecology of the plankton. The filamentous algae are
abundantly represented in spring in the backwaters of the Illinois
River, where they form extensive littoral fringes of "blanket moss,"
which load down the emerging littoral flora. This fringe is fre-
quently stranded by the retreat of flood waters. In some localities,
as in Phelps Lake, it plays a very important part in the food cycle,
since by its decay, as temperatures approach the summer maximum,
it contributes immediately its store of organic nitrogen to the sup-
port of the small algre and flagellates which develop in great num-
bers on those waters at that season. Some species of Spirogyra
and Zygnema have a habit of breaking up into short filaments, and
60
in this condition they have often been taken in some quantity in
the plankton of the river, but they are so plainly adventitious and
irregular that no notice has been taken of them in our enumeration
work, and when possible they have been removed before measure-
ment or deducted by estimation from the volumetric records.
The desmids are few both in species and individuals. Seven
species have been recognized, of which but four are of general
occurrence in the plankton. These are three species of Closterium
and Staurastrum gracile. The latter and Cosmodadium saoconicum
are the only eulimnetic organisms among them. The center of dis-
tribution of the other species is the shore and bottom. The stom-
achs of fish such as the CatostomidcB, the carp, and Dorosoma cepe-
dianum, which often feed upon the bottom ooze or slime about
aquatic plants, usually contain many desmids, including the species
here noted. Other species also are occasionally adventitious in the
plankton, and the list might be considerably extended, though the
absence of extensive peat bogs in the drainage basin of the river
reduces the desmids to a position of much less importance than that
which they occupy in more northerly waters.
As a group they exhibit a well-defined seasonal distribution,
with a vernal pulse at about the time of the volumetric maximum
in April-May and an autumnal pulse of less regular occurrence,
location, and size. The optimum temperature for their appear-
ance in the plankton lies below 70, and in winter months they
occur but rarely.
DISCUSSION OF SPECIES OF CONJUGATE.
Closterium acerosum Ehrbg. Average number, 348. More than
three times as abundant in the previous year. This desmid is
perennial in the plankton, having been found in every month of the
year, but at irregular intervals, and never in large numbers. Its
distribution is such as to suggest that it is at the most only semi-
limnetic in habit. The numbers are too small to follow closely the
seasonal distribution. There are pulses on May 3 (3,200), Septem-
ber 6 (2,400), and November 1 (2,500) in 1898; and in 1897 a pulse
on June 28 (2,000) and one on September 21 (24,000). In previ-
ous years vernal pulses in April and occasional autumnal pulses are
to be noted. In so far as the optimum temperature is indicated, it
61
seems not to lie near either extreme, and above rather than below
the average for the year.
Closterium gracile Breb.* Average number, 49,616 (silk, 305).
This species was found in small numbers from March to December,
and shows pulses on May 17 (1,600) and September ~27"(6,400) at
temperatures of 64 and 73. The tenuity of the form of the
frustule of this species suggests a limnetic habit.
Closterium hmula Ehrbg. Average number, 556. This also is
a perennial species, and is somewhat more abundant and constant
than C. acerosum. It likewise has a vernal pulse, which in 1895
appears on April 29 (2,915) ; in 1896, on May 1 (5,364) ; in 1897, on
May 25 (3,200); and in 1898, on May 24 (6,000). In both this
species and C. acerosum there are slight indications of recurrent
minor pulses which are often coincident in the two species. Nine
such movements appear in 1898. The autumnal pulses are less
regular in their appearance and size than the vernal, and appear
from September to November. The optimum temperatures seem
to lie between 45 and 70. This species is only semi-limnetic, and
never attains the fluctuations which characterize most limnetic
organisms. Doubtless other so-called species of Closterium have
been included among the variable organisms referred here to C.
lumila and C. acerosum.
Cosmarium constrictum Delp. was found occasionally from March
to September, and is probably adventitious.
Cosmocladium saxonicum De By. A single isolated pulse of this
minute limnetic desmid appeared in the filter collections of Septem-
ber, 1897. It was first noted on August 31 and disappeared after
September 29, and was never found at other times in the plankton.
The pulse culminated September 9 at 13,500,000*.
Gonatozygon brebissonii De By. The filaments of this desmid
were noted in the plankton only in March, 1899, attaining a maxi-
mum of 136,800 on the 14th.
Staurastrum gracile Ralfs. Average number, 31. About two
hundred times as abundant in the plankton of 1897. It occurs from
March to January. No vernal pulse was detected, but an autumnal
one of 14,000 appears September 29. It appears in much larger
numbers in the filter-paper collections, and is probably a limnetic
planktont in our waters.
62
Undetermined species of Penium, Arthrodesmus, and Docidium
have been found in the plankton but always singly. They are
doubtless adventitious.
PHANEROGAMIA.
The Lemnacecz are represented in our waters by several species
of Lemna, by Spirodela, and by two species of Wolff ia brasiliensis
and columbiana. The first two genera are predominantly floating
surface-plants, while the last occurs at all levels, is taken with the
plankton, and has been treated in our measurements and enu-
merations as a limnetic organism.
Wolff ia brasiliensis Weddell. Average number, 2 ; in 1897, 13.
It appears irregularly in river planktons from the last of March till
January, and is somewhat more abundant in late summer and
autumn. The seining operations of fishermen in the river and
tributary backwaters have much to do with its appearance in the
plankton of the river.
Wolff ia columbiana Karsten. Average number, 7; in 1897, 41.
With the preceding species. Neither of these species are sufficiently
abundant greatly to affect the ecology or quantity of the plankton
of the river, though they are of more importance in the backwaters.
Owing to their size and duration they compete with the smaller
organisms of the phytoplankton, but do not serve as food for any of
the zooplankton.
PROTOZOA.
Average number, 111,731,000. The number of species exceeds
147 ( + 38), distributed as follows: Mastigophora, 60 ( + 10); Rhizop-
oda,3l ( + 28); Heliozoa,5; Sporozoa (3) ; Ciliata, 45; and Suctoria,
5, the numbers in parentheses indicating the additional forms
whose specific rank was not recognized in the enumerations .
The Protozoa occur in great numbers (Table I.) in every collection
of the year. Owing to the fact that the totals are a conglomerate of
two methods of collecting, of a large number of species of many di-
vergent seasonal tendencies, and of both eulimnetic and adventitious
forms, their seasonal fluctuations have no particular significance which
is not better treated either in connection with the subdivisions of the
class or with the individual species. In the totals, traces appear of
63
the vernal pulse, of the midsummer maximum of the chlorophyll-
bearing Mastigophora, and of the autumnal- winter wave of Ciliata.
The Protozoa, through the Mastigophora, share with the algas
the synthetic function in the elaboration of food from inorganic or
partially disorganized organic contents of the water. _They utilize
decaying organic matter as food, and are thus primary links in the
cycle of food relations. Some of them feed upon bacteria, upon alga3,
or even upon other animals, and thus become secondary or tertiary
links in the chain.
MASTIGOPHORA.
(Plates I. and II.)
Average number, including, without duplication, both silk and
filter-paper collections, 95,856,449. In the collections of 1897
they were five times as abundant as a result, in part at least, of the
extended low-water period, sewage contamination, and extension
of high temperatures during the late autumn of that year (Pt. I.,
PI. XL).
The Mastigophora abound in every collection and occur at all
seasons of the year. Four fifths of them occur, however, between
the first of April and the last of September. They are predominant-
ly chlorophyll-bearing organisms, and have their greatest numbers
during the same season in which the land flora attains its growth.
They spring into abundance with the opening buds of April, and van-
ish from the plankton when frost cuts off the foliage in autumn.
There are, it is true, some species, such as Synura, which grow
luxuriantly at winter temperatures, but these are generally of the
chrysomonad type, with yellowish or brownish chromoplasts. The
bright green chlorophyll-bearing flagellates are in the main summer
planktonts. Since water temperatures do not fall below 32, the
phytoplankton is exempt from this risk of destruction against which
the land flora must provide. We find, accordingly, that the most of the
Mastigophora are wont to occur in diminished numbers and irregu-
larly in the plankton throughout the winter. This appears in the
records of the more common species, and fuller examination would
doubtless greatly increase the number which thus winter over in
reduced numbers.
I have already called attention to the fact that there are in
1898-99 recurrent pulses in the Chl0rophyce& and Bacillariacece at
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66
intervals of several weeks, and that such pulses can also be traced
back into 1897 as far as the collections were made at weekly inter-
vals that is to the early part of July. A similar periodicity on the
part of the Mastigophora the greater part of which are also
chlorophyll-bearing is even more evident. Not only is this
periodicity present in this group, but it coincides approximately in
the location of its maxima and in their relative development with
that found in the Chlorophycece and BacillariacecB. The following
table, which gives the dates of culmination of the pulses of these three
groups from July 1, 1897, to April 1, 1899, will serve to demonstrate
this point more clearly, and a graphic presentation of the data will
be found in Plates I. and II.
There are twenty-two of these recurrent pulses in the period
from July, 1897, to March, 1899. Of the sixty-six possible maxima
only five are missing, or at least not apparent in our data, and but
ten culminate on other dates than the one (of collection) most to be
expected. These ten in every case culminate either a week prior or
subsequent to that in which the other two groups reach their max-
ima. These divergences may be due to the error incident to the
interval of collection, and their approximation in time is still cor-
roborative of the tendency towards recurrent periods of growth.
These exceptions are no greater than might be expected to occur in
the unstable fluviatile environment and within the large margin of
error of the plankton method.
There are twenty-one intervals between July 14, 1897, and
March 14, 1899, with a range in length of 20 to 42 days and an aver-
age of 28.95. The intervals in days with the numbers of instances
of each are as follows: 20 (1), 21 (3), 22 (1), 23 (1), 26 (1),
27 (1), 28 (7), 35 (3), and 42 (3), days. The effect of the weekly
interval of collection is seen in the preponderances at 21, 28, 35, and
perhaps at 42, days. There is evidently a tendency towards the
interval of 28 days. Nine of the 21 pulses are grouped about this
interval; 6, about that of 21 ; while 3 are at 35 and 3 at 42. If there
be such a tendency it is but natural that with a weekly interval of
collection there should also appear minor preponderances at 2 1 and
35 days. Traces of a similar rhythm may be found in the period of
weekly collections in 1896 (Pt. I., Table III.).
In some instances the environmental conditions at these times
of departure are such as to suggest that they may have produced the
67
shifting in the position of the maxima. Thus the pulse of January
25, 1898, appears after a 3 5 -day interval, but in the midst of the
rising winter flood, to whose effect the delay may be attributed.
In both 1896 and 1898 the 28-day rhythm is interrupted at the time
of the vernal pulse in April-May. It appears as though these re-
current pulses -if such exist were submerged in the greater ver-
nal increase. The double summit of the vernal pulse in the curve
of the Bacillariacea and Mastigophora (PI. II.) for 1898 suggests the
compound character of this pulse in the case of these groups of
organisms at least. The time interval in the case of the vernal in-
terruption is also significant. In 1898 there are two pulses between
March 22 and July 19, at intervals of 42 days a total of 84 days,
which is the equivalent in duration of three 28-day intervals.
The total number of species of Mastigophora recorded by me
from the plankton of the Illinois River is over sixty. This number
will be increased to more than seventy if forms not separated in our
enumerations be distinguished as separate species.
The Protomastigina (including the Bicoscecidce and the Cras-
pedomonadida] are well represented in the plankton by passive
limnetic species which are principally sessile on other planktonts.
These are Bicosoeca lacustris, Salpingceca brunnea, S. minuta, and
Diplosiga frequentissima. Asterosiga radiata is a eulimnetic repre-
sentative and Anthophysa vegetans an adventitious one. As a group
they are more abundant during the warmer part of the year.
The Chrysomonadidcz are also well represented, and include the
most abundant flagellates of the plankton of the colder months.
Synura uvella is quantitatively the largest factor furnished by this
group. It is supplemented by Syncrypta volvox, and the various
forms of Dinobryon, Uroglena, and Mallomonas. The last two
genera have more of a summer range of occurrence, but are not of
quantitative importance in the waters of the Illinois.
The CryptomonadidcB are represented only by Chilomonas and
Cryptomonas, and are of somewhat constant, though of minor,
importance quantitatively.
The Euglenidce, on the other hand, are, in our waters at least,
second to no coordinate group in their quantitative importance.
They are individually of relatively large size, and they occur in
great numbers throughout the summer months, replacing the
Chrysomonadida of the colder seasons of the year. Euglena
(6)
68
viridis is the most abundant, and it is associated with other species
of the genus, with species of Amblyophis, Phacus, Lepocindis, Chlo-
ropeltis, Colacium, and Trachelomonas , especially the latter.
The PeridimidcB are quantitatively of considerable importance
in the plankton of our Great Lakes (Kofoid, '95), but in the Illinois
River they are of little significance, at least the larger forms such as
Ceratium. Smaller species such as Peridinium tabulatum and
Glenodinium cinctwn are more abundant. As a group they do not
show any marked seasonal preferences.
The Volvocida, on the other hand, are of more than the usual
consequence in the plankton of the Illinois. The group is repre-
sented by the curious Chloraster gyrans, by the sporadic and meteor-
ic Carteria multifttis, and by the colonial genera Eudorina, Pando-
rina, Pleodorina, Platydorina, and Volvox. As a group they are
almost exclusively summer planktonts.
The Mastigophora as a whole are, next to the BacillariacecB, the
most abundant of the synthetic organisms of the plankton. Their
quantitative importance has not hitherto been sufficiently demon-
strated in the plankton of fresh water, owing it may be to their
escape through the silk net in the ordinary methods of collection.
It seems quite probable also that they may be present in our warm
and fertile waters in much greater abundance than they are in the
colder and clearer waters of most lakes. This is especially true of
the EuglenidcB and Volvocidcz, perhaps less so of the ChrysomonadidcB
and Peridiniidce.
DISCUSSION OF SPECIES OF MASTIGOPHORA.
Amblyophis viridis Ehrbg.* Average number, 63,014 in 1897.
It occurred throughout the summer in 1897, from May to October,
with a maximum of 1,440,000 on August 31. Apparently a sum-
mer planktont but never very abundant.
Anthophysa vegetans (O. F. Mull.) Butschli. This was identi-
fied in the plankton of June, 1898. It is very abundant at times on
various substrata in stagnating water, and from such places becomes
adventitious in detached fragments of colonies in the plankton.
Aster osiga radiata Zach. This interesting colonial and limnetic
choanoflagellate, described originally from the plankton of German
lakes, has been found but a single time in our plankton in the latter
69
part of August, 1896'. It is one of many illustrations of the cosmo-
politan distribution of plankton organisms.
Bicosceca lacustris J. Clark*. Average number, 112,896. Only one
third as abundant in 1896, and four times as many in 1897. This
minute flagellate is found in our waters sessile upon the-filaments of
Melosira, principally M. granulata var. spinosa. It occurs more
frequently upon the dead frustules than upon live ones, and upon
those of the shorter form than upon the longer. It has appeared
also upon Dinobryon sertularia, Pediastrum pertusum, and Richteri-
ella botryoides. It exhibits a considerable range of variation in
proportions, in the amount of lateral compression, and in the length
of the pedicels. These variable forms are, however, connected
with the type as described by Clark, and are not, in my opinion, to
be designated as distinct species. Zacharias ('94) has described
one of these variants as B. oculata. I regard it as a growth condi-
tion of B. lacustris, and not as specifically distinct from it.
Its seasonal distribution in 1898 is somewhat peculiar. It
appears as two quite symmetrical pulses, the first extending from
early in June till the middle of July, and culminating on June 14 at
3,801,600. The approach of this pulse is abrupt and its decline
somewhat gradual. The species does not reappear until September
13. The autumnal pulse culminates October 11 at 486,000, then
gradually declines, and disappears November 1. There is no record
of its occurrence in 1898 outside of these two pulses. In 1897 it is
found irregularly from May to August, and in 1896 in February and
from May to December, with pulses in May, June, July (2) , August,
and October.
In 1898 its optimum temperatures appear at 82 and 65, and its
pulses in other years do not occur below 57. It thus belongs to the
plankton of the warmer months.
Its seasonal distribution falls within that of the limits of its host
Melosira, and in 1896 and 1898 their vernal pulses coincide, and the
same correlation appears in all but one of the pulses of 1896. Not
all Melosira pulses, however, are attended by an increase in Bico-
sceca. Thus in the late summer and fall of 1897 Melosira fluctuated
without any appearance of Bicosceca. In the autumn of 1898 the
pulse of Bicosceca on October 1 1 appears on the decline of the Sep-
tember pulse of Melosira, in which the host made no corresponding
increase. Melosira is thus apparently essential for any marked in-
70
crease of Bicosceca in the plankton, but is not in itself the primary
cause for its appearance in the plankton.
Carteria multifilis (Fres.) Dill.* Average number, 2,365,384.
In 1897 more than one hundred -fold as abundant. This species was
recognized only in the autumnal and hiemal planktons, from
August till January in 1897-98 and from October to February in
1898-99. It is not easily and with certainty identified by the
usual methods of plankton counting, and probably other species of
similar habitus may have been included to some extent ; and, on the
other hand, many Carteria may have been thrown with the "un-
identified" flagellates, especially in earlier years. This species
occurs throughout the whole range of temperatures, and its maxi-
mum development (6,476,400,000) was attained October 5, 1897, at
70. A pulse prior to this appeared September 7, at 2,846,250,000.
From the major pulse in October there is a gradual decline as the
minimum temperatures are reached.
The remarkable outbreak of Carteria in the autumn of 1897 was
associated with unusually low water (Pt. I., PI. XI.) and concen-
tration of sewage and decrease in current. The water of the stream
was of a livid greenish-yellow tinge, due principally to great numbers
of Carteria, which developed to the exclusion or diminution of other
chlorophyll-bearing flagellates such as Euglena, and of diatoms such
as Melosira. This unusual development seems to have been a dis-
turbing factor in the usual seasonal routine of the autumnal plank-
ton of that year.
The distribution of Carteria in the river was remarkable. It
formed great bands or streaks visible near the surface, or masses
which in form simulated cloud effects. The distribution was
plainly uneven, giving a banded or mottled appearance to the
stream. The bands, 10 to 50 meters in width, ran with the channel
or current, and their position and form were plainly influenced by
these factors. No cause was apparent for the mottled regions.
This phenomenon stands in somewhat sharp contrast to the distri-
bution of the usual water-bloom upon the river, which is generally
composed largely of Euglena. This presents a much more uniform
distribution, and unlike the Carteria is plainly visible only when it
is accumulated as a superficial scum or film. Carteria was present
in such quantity that its distribution was evident at lower levels
so far as the turbidity would permit it to be seen. It afforded a
71
striking instance of marked inequalities in distribution within
small areas, of at least one plankton organism.
Carteria showed great variation in the amount of chlorophyll
present. Some individuals were practically colorless. It seems
very probable that in the presence of great abundance^ partially
decayed organic matter such as occurs in a sewage-laden stream,
Carteria may become largely holozoic in its nutrition, as Zumstein
('99) has shown to be the case with Euglena. The literature of
fresh-water plankton contains no record of a similar preponderance
of Carteria in other localities, though its occurrence has been occa-
sionally noted in the plankton.
The chemical conditions under which this great pulse of Carteria
appeared in the autumn of 1897 can be followed in Part I., Plate
XL IV and Table X. The high chlorine and the great increase in free
ammonia and nitrites indicate the decay of sewage; the high
nitrates and albuminoid ammonia show that there was no lack of
some at least of the important sources of food. The two principal
pulses appear September 7 (2,846,250,000) and October 5 (6,476,-
400,000), with a minimum of 680,400,000, on September 21, sep-
arating them. Both of these pulses are attended by sharp declines
in nitrates and nitrites and free ammonia, and very slight decreases
in organic nitrogen and albuminoid ammonia. Either the first
three substances named or those matters which supply them by
their decay, are thus noticeably utilized at the times of these pulses.
The relation of the Carteria to the volumetric pulses is (Pt. I.,
PI. XI.) not a constant one. The Carteria pulse of September 7
lies in a slight depression between two maxima of the volumetric
curve, and a week prior to the autumnal culmination on September
14 at 19.8 cm. 3 per m. 3 . It thus appears during the growth period
of this volumetric maximum. The second and larger pulse of
Carteria, on October 5, coincides with the second volumetric maxi-
mum, and in fact fluctuates throughout with it. Though Carteria
constitutes but a small part of the actual catch of the silk net,
owing to leakage through the silk, it is apparently an important
factor in the food cycle which builds up such maxima.
Ceratium brevicorne Hempel. This species appeared in small
numbers in isolated instances from April through October. It
varies towards C. hirundinella, but the small numbers in which it
has occurred have not as yet afforded sufficient ground for regard-
72
ing it as a variety of that species. It occurs most frequently in
August and September, and is apparently a warm-water planktont.
Ceratium cornutwn Ehrbg. was found but once in June, 1896.
Ceratium hirundinella O. F. Mull, was not noted in our plank-
ton in 1898, but in 1896 was found from June to October, with a
pulse of 19,200 on June 6. It was recorded only at temperatures
above 57, and is apparently a warm-water planktont. It has but
an insignificant part in the potamoplankton of the Illinois River
and its backwaters, though quite abundant in the summer plank-
ton of Lake Michigan (Kofoid, '95). It seems not to have survived
the transit through the sewage-laden waters of Chicago River or to
thrive in the conditions prevailing in the Illinois River, though
common generally in fresh- water plankton of the temperate zone.
Chilomonas paramcecium Ehrbg.* Average number, 555,000.
This flagellate, which is frequently abundant in aquaria or stag-
nant water, appears also in the plankton of the Illinois River.
There is in 1898 a vernal pulse, culminating at 10,800,000 on April
26, and there are scattered records from October to February.
Chloraster gyrans Ehrbg. This rare and unique flagellate was
found in but two collections in July and August, 1898 and only
in small numbers.
Chloropeltis monilata Stokes.* Average number, 362,941 in
1897. This is a summer planktont, appearing at irregular inter-
vals from the last of May until the middle of September. It was
not found in 1898. A maximum of 10,800,000 appears on August
31.
Colacium calvum Stein. The attached stage only of this flagel-
late was observed , and was recorded only in 1 8 9 6 and 1897. It appears
from the middle of April to the first of October, and is usually
found upon Polyarthra platyptera. It has occurred occasionally
upon several species of Brachionus and upon Chydorus sphczricus.
The largest number recorded (162,792) appeared on April 17, 1896,
upon Polyarthra, usually upon the body and more rarely upon the
oar-like appendages. It is often exceedingly abundant upon the
planktonts of backwater ponds.
Colacium vesiculosum Ehrbg. This species is much less abun-
dant than the preceding species in our waters, and was found only
in June and September, upon Cyclops albidus and Polyarthra.
73
Cryptomonas ovata Ehrbg.* Average number, 121,154. This
species has been recorded principally in the autumnal or hiemal
plankton. It escapes through the silk net readily, and was rarely
found in collections of earlier years. In 1895 it occurred from July
till the last of October, and in 1898 was common in the December
plankton.
Dinobryon sertularia Ehrbg. Like most typical planktonts,
Dinobryon is an exceedingly variable organism, and the varia-
tion finds its expression in the form and proportions of the
loricas and in their arrangement and continuity in colonies.
Divergences from described and figured species are thus at once
apparent, and they have been utilized by systematists, notably by
Lemmermann ('00) and by Brunnthaler ('01) as the basis for the
establishment of a large number of new species. The validity of
these species, in my opinion, must rest ultimately upon careful
experimental evidence of their present mutual genetic independence
under normal conditions of growth. From my own observations
upon large numbers of colonies and individuals distributed through-
out the range of their seasonal recurrence in six years in our waters,
I am inclined to regard all as belonging to a single species, and the
different types as mere growth varieties. The rapidity of growth
and the age of the individual or of the colony are, I believe, impor-
tant factors in the determination of the form of the lorica, and its
various forms are therefore not of specific value, but rather of
physiological significance. It is a simple matter to find individuals,
or even colonies, conforming to the descriptions of the several
species, but it is not so easy to refer all individuals and all colonies
to the described types. They intergrade nay, more, two, or even
more, "species" are not infrequently combined in the same colony.
I have never found all the forms in a single colony, but such com-
binations as angulatum-divergens, diver gens-angulatum-stipitatum,
sertularia-angulatum, and sertularia-undulatum have been observed
by me. These combinations are most frequent in large colonies,
and, indeed, the number of "species" in a colony is apparently a
function of its size. The slender growing tips are wont to assume
the stipitatum type of lorica and colony, and the older loricas at the
base to conform to that of sertularia, divergens, or angulatum.
Small colonies as a rule belong to a single "species." These com-
binations are generally most evident during the maximum period
74
of growth; that is, when Dinobryon is multiplying rapidly, though
they may appear at any season of its occurrence.
In the enumeration of Dinobryon five types were recognized, and
the individuals were assorted to these "species," viz. : D. sertularia,
stipitatum, diver gens, angulatum, and undulatum. Some corrobora-
tion of the view that we are dealing with a single variable organism
and not with five distinct species may be seen in the coincidence of
the seasonal distribution, and of the rise, culmination, and decline
of the pulses of the five different forms.
Since these varieties have such a similar seasonal distribution I
shall treat them as a whole, discussing subsequently any individual
peculiarities which are noteworthy. The average number of
individuals of Dinobryon sertularia, including all its varieties, in
1898 was 1,979,785. In 1897 the average was much smaller
(79,352) owing to the few collections in the winter, when it is most
abundant, and to its suppression in the prolonged low water of the
autumn of that year. The relative frequency of these different
varieties for I shall treat them as such is shown by the average
per cubic meter for the year in 1898, viz.: D. sertularia, 407,602;
D. sertularia var. stipitatum, 603,911; D. sertularia var. divergens,
866,083; D. sertularia var. angulatum, 101,358; D. sertularia var.
undulatum, 831. These figures are only approximate, since colonies
containing more than one variety have all been included with the
predominant variety in the colony, which is usually sertularia or
divergens, consequently angulatum and undulatum are more
numerous than indicated by these figures.
The seasonal distribution of Dinobryon in our waters is well
defined, and is sharply limited to the period from November to June.
Its earliest recorded appearance was November 8 in 1898, while in
1896 and 1897 it was not found until in December. It lingers well
into June in 1896 and 1898 the two years in which the spring
collections were of sufficient frequency to trace its decline. In 1898
the latest record was on June 28. Most of the records after May
are irregular and sporadic. It is thus absent from the plankton of
the Illinois River from the last of June till November or December.
In 1895-1896 there was also a winter interval in which no Dino-
bryon was recorded during the December- January flood (Pt. I., PI.
IX. and X.). In 1897-1898 a similar interval appears, and con-
tinues almost to the end of the slow rise of the flood which culmi-
75
nated in March. Rising floods thus do not favor the development
of Dinobryon in channel waters of the Illinois.
The interval of collection in 1894-95 is too great to trace the
seasonal fluctuations of Dinobryon, though there are indications of
a maximum pulse on April 29. In 1895-96 there is^ a slight de-
velopment in November prior to the rise of December, in which
Dinobryon again disappears. A slight pulse of 3,192 appears on the
declining flood (Pt. I., PI. X.) on January 25, and declines again
with the rise in February to reappear on February 20 at 42,588.
Another decline in Dinobryon attends the rise in river levels in
February-March, and after a fortnight of falling levels a third
pulse of 2,531,280 is seen on March 17. Two other pulses attend
the decline of this flood, one upon April 29 (800,064) and the other
on May 18 (339,624). On the decline of the June rise of this year a
late and unusually large pulse for the season appears (June 11) at
2,438,400. An examination of the hydrograph will indicate that
almost without exception these pulses attend the run-off of im-
pounded backwaters after recent invasion, or, as on April 29 and
May 18, after a temporary check in the run-off. During those
times when the channel contributes to the backwaters, that is, dur-
ing rising floods, Dinobryon declines in numbers ; and, on the other
hand, it reaches its greatest development in channel waters during
the run-off of the flood.
In 1896-1897 the interval of collection (Pt. I., Table III.) is
again too great to trace satisfactorily the fluctuations of Dinobryon.
There is a pulse on December 3 of 157,609 and on April 27 of
172,800.
In 1897-98 Dinobryon appears first on December 7, with a pulse
of 1,807, 200, during a period of low water and ice blockade with no
backwater contributions. It declines, and after December 21 does
not again return until March 22, when an isolated record appears.
The vernal pulse begins April 19 and culminates May 10 at 84,-
841,600 on the declining spring flood (Pt. I., PL XII.). Dinobryon
declines at once during a fortnight of rising water, and two minor
pulses on the decline of the flood one on June 7 of 70,400 and one
on June 28 of 219,840 complete its vernal cycle.
The hydrographic conditions in 1898-99 were very different from
those of the preceding season, and we find a marked change in the
seasonal occurrence of Dinobryon. From November to March
76
there are three rises to overflow stages (Pt. I., PL XII. and XIII.)
with intervening declines of a month's duration. There is a pulse
of Dinobryon in each of these periods of declining flood. The pulse
of 275,200 on December 20 follows the November flood, and it is
followed by a minimum of 1,500 on the rising flood of January 10.
The numbers slowly increase until a meteoric rise on February 7
to 6,486,700 and on February 14 to 22,621,440 is followed again by
another decline, to 25,920 on February 28, with the sudden flood of
that week. During the .maximum flood stage in March (Pt. I., PL
XIII.) no Dinobryon was recorded, but it reappeared again on
March 21. The suspension of our plankton operations interrupted
the further tracing of the fluctuations.
From the facts above detailed it is very evident that the pulses
of Dinobryon occur in channel waters at times when the run-off of
impounded backwaters is making its greatest contribution to the
river plankton. These are times of greatest stability of the en-
vironment in all respects save river level and its sequences. The
impounded waters have come from regions of slight current and
decaying vegetation, and there has been time in those localities for
the decay of sewage and debris, and for the growth of planktonts
such as Dinobryon. These conditions of the environment are
therefore favorable for the growth pulses of Dinobryon. The
phenomenon of pulses of growth is not, however, to be considered
as merely the result of declining floods. These afford a favorable
environment and doubtless determine within certain limits the
time and the extent of the pulse. The phenomenon is one common
to most plankton organisms, and occurs in Dinobryon of lakes where
floods are of little significance.
Any evidence of recurrent minor pulses in Dinobryon at brief
intervals is lacking.
Dinobryon has been found in our plankton through practically
the whole range of temperatures, but it disappears when maximum
summer heat is reached and does not return until the water cools
to 45 or lower. Large pulses, such as that of February 21, 1899
(22,621,440), have developed at temperatures approximating 32,
and largely under the ice. The vernal pulse of April-May has been
recorded at temperatures ranging from 60 to 79, but generally
nearer the former. No well-defined optimum temperature appears,
and the seasonal distribution suggests that the high temperatures
77
of our summer waters are inimical to Dinobryon. That its absence
from the plankton at that time is not due merely to low- water con-
ditions is shown by the December pulse in 189 7, under the most pro-
nounced type of such conditions.
Dinobryon is a common planktont in the Great-Xakes (Kofoid,
'95) during the summer months, but surface temperatures here
rarely exceed 68, and are 10 to 20 below those of the Illinois
River. In German lakes Apstein ('96) finds the maximum develop-
ment of Dinobryon in June and a continuance through the summer
in reduced numbers, but temperatures are also 10 to 20 (F.)
lower than in our waters. In the case of D. stipitatum there is a
second maximum in August. Lauterborn ('93) finds Dinobryon
throughout the winter in the plankton of the Rhine, with a maxi-
mum in April-May, with diminished numbers during the summer,
and a second maximum in September.
The filter-paper collections give very much larger numbers,
owing partly to the inclusion of small colonies which escape
through the meshes of the silk net in the usual method of collec-
tion. The numbers are increased at least thirty-fold if filter col-
lections are utilized instead of silk, as above.
The size of the colonies in the collections varies greatly, the
averages ranging from three to forty-eight cells. The maximum
pulse is attended or followed by a considerable decrease in the size
of the colony. In the pulse of February 21, 1899, the average num-
ber of cells in the colony falls from thirteen to sixteen, during the
rise of the pulse, to seven, at its culmination. On the pulse of
May 10, 1898, the average is thirteen, and a week later, when the
pulse declines from 16,153,600 to 43,200, the average size of the
colony drops to three cells. Cysts also are most frequent during
and subsequent to maximum development. Dinobryon is some-
times covered with large numbers of minute choanoflagellates,
probably Salpingosca minuta Kent. Frequently colonies occur in
which only the younger cells are alive.
Dinobryon is, in the light of its distribution, one of the impor-
tant synthetic planktonts of the colder months, and is one of the
primal links in the chain of food relations of that season, serving
as food for some of the winter Cladocera and Copepoda. The fact
that its maxima frequently occur when volumetric minima appear
as, for example, on February 21, 1899 indicates that Dinobryon
78
does not directly contribute much, even at its maximum develop-
ment, to the volume of the plankton taken in the silk net. On
the other hand, its rapid multiplication, as evidenced by its meteoric
pulses, may serve to build up a more permanent and bulkier animal
plankton, and thus indirectly, in a cumulative way, it may be of
considerable quantitative importance.
The inclusion of all the variants of Dinobryon as a single species
has been favored by Wesenberg-Lund ('00), who regards D. stipi-
tatum as the summer form of D. sertularia. In our plankton, D.
stipitatum has occurred sporadically in December and March, but
it is most abundant during the vernal pulse in April-May. Its
distribution thus in the main supports that author's contention in
that it is found during the warmer portion of the seasonal cycle of
Dinobryon in our waters, though not in our summer plankton. It is
not desirable in this connection to enter further into a discussion of
problems which have been raised by the splitting up of Dinobryon
into so large a number of forms. Lemmermann has found seven-
teen species and varieties within the limits of the subgenus Eudino-
bryon. A discussion of their validity involves not only some per-
plexing problems of synonymy, but also an extensive examination
of a large amount of material showing seasonal changes, and,
above all, a series of experiments which shall demonstrate the
limits of variation within a known line of descent and in the sea-
sonal range of environmental conditions. It involves, moreover,
the fundamental question of the criterion of species. The papers
of Lemmermann ('00) and Brunnthaler ('01) have appeared since
my work of enumeration was completed. I recognize among the
forms which they have sought to establish the following which
occur in our plankton: D. sertularia Ehrbg., D. sertularia var.
thyrsoideum (Chodat) Lemm., D. sertularia var. alpinum Imhof,
D. protuberans Lemm., D. sociale Ehrbg., D. stipitatum Stein, D.
stipitatum var. americanum Brunn., D. stipitatum var. bavaricum
(Imhof) Zach., D. elongatum Imhof, D. elongatum var. undulatum
Lemm., D. cylindricum Imhof, D. cylindricum var. palustre Lemm.,
D. cylindricum var. schauinslandii (Lemm.) Lemm., D. cylindri-
cum var. ped^forme (Lemm.) Lemm., D. cylindricum var. diver gens
(Imhof) Lemm., andD. cylindricum var. angulatum (Seligo) Lemm.
As a result of my attempts to refer all of the individuals which I
have seen in my work of enumeration to species, I am of the opinion
79
that we are dealing in the case of the species of Dinobryon above
cited with a single variable organism, whose extremes of variation
only have been regarded as separate species. The connecting links
are sufficiently abundant still and the union of several types in a
single colony is sufficiently frequent to lend some weight to my con-
clusions with regard to those forms which have been under my
observation. In the interests of utility as well as in the interests
of well-grounded taxonomy, it is extremely desirable that the
establishment of new species among variable plankton organ-
isms should be attempted with extreme caution and only after the
fullest study of the range and conditions of variability. The insta-
bility of the taxonomic structures which Brunnthaler and Lemmer-
mann have recently raised, is evidenced by the differences in syno-
nymic, varietal, and specific rank given to the variants of Dino-
bryon by these two systematists, who have but recently mono-
graphed the group, largely if not wholly from the systematic point
of view. The changing estimate of validity which Lemmermann
himself has put upon his own species or varieties for example,
schauinslandii, pediforme, and curuatum rgives further evidence
that the basis upon which they rest is at the best but slight.
It is my firm conviction that the establishment of new species
among the organisms of the plankton of fresh water can be
satisfactorily accomplished only after careful analysis of the limits
of variation within the range of environmental conditions. Stand-
ards less comprehensive than this can yield results of but temporary
or local value and can lead to but little permanent advance in
science, and they bring only perplexity and chaos where order
should reign.
Diplosiga frequentissima Zach.* Average number, 1,736,538.
This minute flagellate is found upon the rays of the colonial diatom
Asterionella, often in great numbers and so thickly set as to leave
little unoccupied space. It was found in each year at the time of
the vernal pulse of Asterionella in April-May, and was as a rule most
abundant immediately after the maximum growth of Asterionella
had been attained. Beyond an isolated occurrence in January it
was not recorded at other times than during the months of April
and May.
Eudorina elegans Ehrbg. Average number, 14,362. About
twice as abundant in 1897. The distribution of this species is
80
somewhat erratic. It has occurred in every month from February
through October, but in smaller numbers and sporadically in the
colder months. In 1898 its seasonal curve is of characteristic form.
It makes its appearance March 1 5 , and is continuously present until
the end of September. There is a vernal maximum April 26 of
240,000, but no corresponding autumnal one. In 1898 there are
indications of recurrent pulses at brief intervals which coincide in
location immediately or approximately with similar ones of Gonium
and Pandorina. These pulses occur March IS (3,600), April 5
(2,800), April 26 (240,000), June 14 (60,000), August 2 (8,000),
August 23 (3,200), and September 20 (2,000). The minima
between these pulses in all cases but one fall below 1,000. In
1897 a vernal pulse was not detected, a maximum of 496,000
occurring August 31, and but three mindr pulses appearing. In
1896 this species appeared in the plankton on February 20,
and remained until the end of August with a month's interrup-
tion in May- June. There were no marked pulses, exceeding
15,000, in that year. The absence of the spring flood (Pt. I.,
PI. X.) and the disturbed hydrograph of the summer may account
for this suppression of development in Eudorina. The distribu-
tion in preceding years is also irregular.
Eudorina begins its seasonal development at temperatures but
slightly above 32, but any considerable growth is not attained
until at least 45 has been reached, and the largest pulses on record
have been at the close of the period of maximum summer heat at a
temperature of 80, and the vernal pulses have been at 60 or above.
The disappearance of Eudorina from the plankton in the early fall,
about the tim that foliage is killed by autumnal frosts, has been
constant in the different years.
Eudorina is not sufficiently abundant to be of any considerable
importance in determining directly the volume of the plankton.
It serves as food for many of the rotifers, and is itself frequently
parasitized by Dangeardia mammillata Schroder, which destroys the
cells but leaves the matrix intact. There are times when it is
hardly possible to find perfect colonies, and when it is not unusual
to see colonies swimming about propelled by one or two surviving
cells.
Euglena acus Ehrbg.* Average number, 214,807. Found from
the middle of March till the first of November, and most abundantly
81
in late summer and early autumn. It escapes through the silk net
readily, and no marked pulses in occurrence appear in the erratic
data of the filter-paper collections. It is found in the water-bloom,
and is predominantly a warm- water planktont.
Euglena deses Ehrbg. Occurs occasionally in the plankton and
water-bloom during summer months.
Euglena elongate Schew.* Average number in 1897, 278,970.
It is found irregularly in our plankton and water-bloom from July
to October. Originally described from New Zealand.
Euglena oxyuris Schmarda.* Average number, 960,769. Next
to E. viridis this is the most abundant member of the genus in our
plankton. It is abundant during the summer, especially towards
its close during low-water conditions, when the water-bloom,
in whose formation it shares, is best developed. There is no
vernal development, and the fluctuations are but slight in com-
parison with those of most organisms of the plankton: There is a
slight indication of recurrent pulses at intervals of a few weeks.
Its optimum temperature lies near that of maximum summer
heat, that is, about 80, though some tendency to run over into
autumn months is manifest.
Euglena sanguinea Ehrbg. There are only sporadic occurrences
of this species in the plankton. It is found along with E. vindis
among matted growths of Lemnacea, and on exposed and reeking
mud flats, where it forms patches of bright red color often of large
extent. It may be only a physiological condition of E. viridis,
with which it is always found. It has appeared in the plankton
most frequently in September, though found elsewhere throughout
the summer.
Euglena spirogyra Ehrbg. Found but once in October, in the
river plankton.
Euglena viridis Ehrbg.* Average number, 1,571,731 ; from silk
collections only 8,653. This is the most abundant of the larger
green flagellates in our plankton, and constitutes the greater part of
the water-bloom of summer months, when it forms towards four
p. m. a livid green scum on the immediate surface of the water.
Collections of the silk net give no clue to its abundance and shed no
light on its seasonal distribution. The filter-paper collections indi-
cate its presence from March to December, but in numbers only
during the warmer period, from May to October. There is no ver-
82
nal pulse though there are slight traces of minor irregularities, and
on September 7, 1897, a single unusual deyelopment of 58,000,000.
Its optimum temperatures lie close to the maximum heat of summer
months. It is found not only in water-bloom and plankton, but
also along shores, on mud banks, and in sequestered pools and bays
where temperatures reach 90 and over. Lightly colored and semi-
transparent individuals of this and other species of the genus are
found frequently in the plankton, suggesting an approach to holo-
zoic nutrition in nature, such as Zumstein ('99) has demonstrated
experimentally in E. gracilis. Euglena is quantitatively one of the
most important links in the chain of food relations of the summer
plankton, converting nutrient matters in the water, both organic
and inorganic, into food for the Rotifer a and Entomostraca of that
season of the year. It in a measure replaces the diatoms, some of
which decrease in number or disappear during the warmer months.
Glenodinium cinctwn Ehrbg.* Average number, 1,360,192.
This species is generally present from the middle of March till the
end of September, though sporadic occurrences are found in winter
months. There is a pulse on March 29 of 4,260,000 at a temperature
of 49, and another August 9 of 25,200,000 at 83. This small
planktont usually escapes through the silk net. It may be that
several species have been included, as the conditions of plankton
enumeration do not permit close scrutiny of such small organisms,
lacking prominent structural characteristics. It seems to be a
perennial planktont with a wide range of temperature adaptation,
and with a growing period approximating that of the land flora of
our latitude.
Gonium pectorals O. F. Mull. This colonial flagellate has been
found in the water-bloom in large numbers, especially in the back-
waters. It was taken in the river plankton in 1897 and 1898 in
May and again in August and September. These pulses coincide in
location with those of Pandorvna and Eudonna.
Lepocinclis ovum Ehrbg.* Average number, 401,538; silk
3,719. This species appears in the plankton in April and continues
until the end of October, with sporadic appearances in winter
months. There is no vernal pulse, and in both 1897 and 1898 maxi-
mum numbers, 43,200 and 50,400, occur at the height of midsummer
heat in August. In both years there are well-defined recurrent
pulses at intervals of three to six weeks to be traced in the silk
83
collections. The optimum temperatures plainly lie near the maxi-
mum, that is, about 80, and the season of growth approximates
that of the land flora, being limited to the months of April-Septem-
ber. This is a variable organism, and a number of species have been
described in the genus in recent years. Many of these" occur in our
waters, but no attempt has been made to separate them, since they
are based on minute characters.
Mallomonas plosslii Perty. and M. producta Zach. These two
forms will be treated together, as in my opinion they are merely
divergent variants perhaps seasonal of a single species. In 1898
M. plosslii was found but three times in June and July and M.
producta eight times from May through September. In 1897 the
latter only was recorded, and in September and October. In 1896
M. plosslii appeared in July and M. producta in April and August.
In 1895 M. producta alone was recorded, and that in November.
The data are hardly sufficient for generalization, but so far as they
go they indicate that producta is more prevalent in late summer and
autumn and plosslii in early summer, the more attenuate form
(producta} in the warmer season.
Biitschli ('80-' 89) has intimated that there may be some genetic
connection between Mallomonas and Synura uvella. Certain
features of its occurrence in our plankton lend their support to this
view. Synura in our waters is a winter planktont, with December
and February or March pulses. Mallomonas is a summer planktont,
making its first appearance during the time of the decline of Synura,
and when many of the colonies of the latter are breaking up into
their individual zooids. Again, the differences in structure and size
between the two genera are quite superficial, and might result from
the growth attending the free life of a Synura zooid and its prepara-
tion for sporulation. It is a noticeable phenomenon that the pro-
portion of sporulating individuals of Mallomonas in the plankton is
exceptionally large among all plankton organisms. "Free cells"
of Synura are plainly referable to that genus by their resemblance,
and by the fact that they are often united in clusters of several indi-
viduals forming fragments of disintegrating colonies. It may be
that some reproductive phase, as conjugation, intervenes between
the free-cell condition of Synura and the Mallomonas stage, and
that the relatively smaller numbers of the latter are due to the in-
frequency of this process. While the features of seasonal distribu-
(7)
84
tion, structure, and sporulation thus suggest the possibility that
Mallomonas is a free zooid stage leading to sporulation in Synura,
they do not demonstrate it, and the genera must stand in statu quo
until breeding experiments shall clearly demonstrate the full life-
cycle of Synura.
Pandorina morum Bory. Average number, 6,957. In 1898 this
organism was about half as abundant as Eudorina, but in 1897 it
more than equals it. On account of the small size and the motility
of the colonies many of them escape through the silk, so that it is
not so adequately represented in silk-net collections as Eudorina.
It is probably the most important quantitatively of the Volvocidcz
in our plankton. It occurs from April to October, with a few spo-
radic appearances in March and up to January. Its greatest growth
occurs from May to October. There is no predominant vernal pulse
in 1898, but a series of smaller ones culminating May 3 (48,400),
June 14 (60,000), July 26 (63,200), and August 30 (3,200), all upon
declining floods (Pt. I., PL XII.) and coincident with pulses of
other VolvocidcB Eudorina and Gonium. In 1897 its seasonal dis-
tribution was also similar to that of these genera, exhibiting a max-
imum pulse August 31 of 638,000 at 80. In 1896, a year of inter-
rupted hydrograph (Pt. I., PL X.), Pandorina attained no marked
development. Its optimum temperatures lie at and above 60,
and its larger pulses appear during the season of maximum temper-
ature, that is, at about 80. Pandorina does not attain any marked
autumnal growth, but declines in September, and as a rule dis-
appears in October. The period of its growth thus lies within that
of the land flora.
As in Eudorina, so also here, parasitism by Dangeardia mammil-
lata is of frequent occurrence. Pandorina is an important element
in the food of summer rotifers such as Brachionus.
Peridinium tabulatuni Ehrbg.* Average number, 3,875,769;
silk, 3,711. This is a perennial planktont, having been found in
every month of the year. Its principal development is, however,
reached during warmer months, from May till September. In 1897
the maximum pulse of 172,800 was on August 10, and in 1898 one of
66,800 fell on July 26, the temperatures being 81 and 89 respectively.
The only exception to this predominance in warm months is an iso-
lated pulse of 2,400 which developed on the declining flood of Febru-
ary, 1899 (Pt. I., PL XIII). The absence of any autumnal development
85
of this species is noticeable. Its optimum temperatures lie close to
the summer maximum (80), and though perennial, its occurrences
at other seasons than late spring and summer are irregular and its
numbers few. Its seasonal distribution in German- lakes, as re-
ported by Apstein ('96), is similar to that in the Illinois River.
The Peridiniidas play but an insignificant part in the plankton of the
Illinois River.
Phacus longicaudus Ehrbg.* Average number, 61,153; silk, 3 ,03 1 .
This species in 1898 made its first appearance in the plankton on March
23 and continued till November 15. The species is small enough to
escape through the silk net, and the data from such collections do
not fully express its seasonal fluctuations. There is no marked
vernal pulse, and there are traces of but a few small ones during the
summer, the largest in 1898 being one of 35,200 on September 27.
The distribution in previous years is much the same. A well-sus-
tained development throughout the warmer months save when
rising floods, as that of May, 1898, reduce the numbers indicates
that the optimum temperature for the species approaches the
summer maximum (80). There are almost no occurrences below
45. This is the most abundant member of the genus in our plank-
ton, but it is not quantitatively an important element therein.
Phacus pleur one ctes Nitzsch.* Average number, 450,000; silk,
298. It is less abundant (from one fifth to one tenth) than P.
longicauda in the catches of the silk net but apparently much more
abundant in the filter-paper collections, which may be due in part
to its smaller size and greater tendency to escape through the silk
in the collections of the net. Its occurrences are even more closely
limited to summer months from June till September. There is no
vernal development, and the largest numbers occur during the
period of maximum heat. Pulses are but feebly defined. It is also
a summer planktont.
Phacus pyrum Ehrbg. was found but once on August 10, 1897.
Phacus triqueter Ehrbg. occurred in small numbers during July
and August, 1897.
Platydorina caudata Kofoid. Average number, 17. In 1898
this interesting new genus of the Volvo cidoe was found in the plank-
ton only in the latter part of July. In 1897 it was much more abun-
dant (average number, 21,963) and ranged from July 14 to October
86
12. There was a pulse on July 21 of 18,400 and another on Septem-
ber 7 of 600,000. In previous years the occurrences were scattering,
but confined to July, August, and early September. It is evidently
a summer planktont, whose optimum temperature lies near the max-
imum attained by our waters. No record of occurrence below 60
was made. The smaller and younger colonies escape readily
through the silk net. Its pulses in 1897 coincide very closely with
those of Gonium, Pandorina, Eudorina, and Pleodorina.
Pleodorina calif arnica Shaw. Average number, 11. In 1897
this species, in common with other members of the family, was much
more abundant than in any other year of our work, stable conditions
of low water with the accompanying sewage contamination seeming
to favor its development. The earliest record for P. calif ornica
in the plankton is May 18, 1896, at 71. This was a year of low^er
water and higher temperatures than usual in spring months (Pt. I.,
PI. X.). In other years P. calif ornica did not appear until June or
July. It continues into September, the latest record in 1895 being
October 2. In 1897 there were pulses on July 21 (5,600) and
September 7 (4,000). The occurrences at other seasons are too
scattered to trace the seasonal fluctuations, but there is a well-de-
fined predominance during the period of maximum heat. This is
evidently a summer planktont, whose optimum temperature lies
near 80.
Pleodorina illinoisensis Kofoid. Average number, 6,91 7 in 1897.
This is somewhat more numerous than the preceding species, and
its range of occurrences is quite similar. Its maximum pulse in
1897 (180,000) is on August 31, a week earlier than in other members
of the family. These pulses of the VolvocidcB occur (Pt. I., PI.
XLIV.) in a depression of nitrates and just prior to the volumetric
pulse of September, 1897. This pulse is doubtless built up partly
at their expense. Their decline in numbers corresponds with its
rise. This is also a summer planktont, and was not recorded
below 71.
Salping&ca brunnea Stokes.* This species was not recorded in
1898. Average number in 1897, 1,887,356. It occurred on May
25 and July 21, dates of culmination of pulses of Melosira granulata
var. spinosa. In August-September a pulse occurs, culminating
September 7 at 47,250,000 a week after the culmination of a Mel-
osira pulse. In 1896 (silk collections only) it was present through-
87
out most of the summer, attending only approximately the sup-
pressed and interrupted pulses of Melosira in that year of disturbed
hydrograph. It has been recorded from the latter part of April
till the middle of September, and, as a rule, above 60. This beau-
tiful little choano flagellate is sessile upon the filaments of Melosira,
principally upon the variety spinosa, and but rarely upon M. varians
or other planktonts such as Pediastrum. It is often associated
with Bicosceca lacustris and is usually found upon the sides of the
filaments, the bowl of the transparent brownish lorica being closely
sessile upon the diatom. In one instance a lorica was found upon
the corner at the end of the filament. The lorica had adapted
itself to this novel situation by an angular indentation fitted upon
the corner of the diatom.
Syncrypta volvox Ehrbg. Average number, 625. This species
has a definite and somewhat unusual seasonal distribution. In
1898 it was found from March 1 to April 12, and reappeared Novem-
ber 8, attaining a maximum of 13,500 on December 6, and of 43,000
on January 1, declining then to 800 and rising on February 14 to
4,800, and subsequently disappearing in the flood waters of March.
It was not recorded in 1897. In 1895 it appeared September 27 and
continued for a month, reappearing in February and March, and
not occurring after April 10. It has attained its largest develop-
'ment at minimum temperatures under the ice 43,000 January 3,
1899, at 32.7. The greater part of its occurrences in 1898-1899
lie very near this temperature, and but three in all the years lie
above 50. It is par excellence a winter planktont, or at least a
cold-water one.
Its occurrences in 1895-1896 lie near the beginning and the
close of the seasonal pulse of Synura. In 1898-1899 the pulses of
Syncrypta coincide in location with or immediately follow those of
Synura. The resemblance of Syncrypta to small colonies of
Synura is striking, and this fact combined with the relation of their
seasonal fluctuations raises the query if Syncrypta may not be an
encysting stage of the Synura colony. Its life history should be
fully worked out.
Synura uvella Ehrbg. Average number of colonies, 8,463.
The seasonal distribution of this chrysomonad flagellate is some-
what similar to that of its near relative Syncrypta. It is a
perennial, though predominantly cold-water, planktont. It appears
88
in the December plankton of 1894, but was exterminated from the
channel plankton taken in the following February by the stagna-
tion attending the long-continued ice blockade. It reappears in
April, and again disappears promptly, but does not return until
September 12, and not in numbers until October. There are pulses
November 20 (506,800) at 42.8, and December 30 (362,520) at 36.5.
The December pulse is followed by a decline, with a rise during
February to a well-sustained maximum during March, approaching
400,000, and at from 35 to 48. The decline follows in April, and
there are only isolated occurrences in small numbers at irregular
intervals during the summer. Continuous occurrence begins again
in September, and numbers rise rapidly in October. There is a pulse
of 542,699 on December 3 at 32.2, and another on March 22, 1897,
of 159,500 at 43.8. Synura is very rare indeed in the summer of
1897, and in the prolonged low water, sewage contamination, and
higher temperatures of the unusual autumn of that year it does
not reappear continuously until October 26, at 59, and does not
exceed 1,000 until December 7, at 32. There is a low maximum
of 98,700 on December 14 at 36, followed by a decline during the
rising flood of January-March, 1898. The slight cessations in the
flood invasion (Pt. I., PL XII.) in January and in the second weeks
of February and March produce prompt responses in immediate
rise in numbers in Synura. Finally, a low maximum of 320,600 is
attained upon the crest of the March flood, on the 29th, at 49.
This is followed by a decline during April and a few scattered
appearances during the summer. Synura returns at the end of
October and rapidly mounts to a pulse of 1,999,500 on November
29 at 35 with the first decline of the November overflow (Pt. I.,
PI. XII.). A second pulse of 2,764,800 on December 20 at 33,
under the ice, gives way to a decline to 51,600 towards the end of
January, 1899, during rising water. On February 14 another pulse
(348,800) appears at 32.5, under heavy ice, and declines again in
the sudden flood of the last days of February, but recovers quickly
with a maximum pulse of 898,800 on March 7 at 32.8. Within a
fortnight this falls to the low level of 9,800, but its further history
was not followed.
From these data it is evident that in our waters at least Synura
is limited to the months from October to April, except isolated and
irregular occurrences of small numbers during the summer. Its
89
optimum temperatures lie below 50, and its greatest development
has taken place in minimum temperatures under the ice. Rising
floods and disturbed hydrographies conditions tend to reduce its
mimbers or to suppress its development, while declining floods
initiate increase in numbers and favor the appearance of pulses.
A "late" autumn delays the appearance of Synura.
Not only are colonies of Synura found in the collections, but at
times large numbers of free cells make their appearance. These
are released by the breaking up of colonies, and occur in all degrees
of isolation. It seems to be a natural phenomenon, and occurs
most abundantly with or immediately after the crest of the pulse.
Thus the pulse of December 29 (1,999,500 colonies) was attended
by 21,600,000* free cells on that date. A week later there were
1,693,500 colonies and 57,600,000 free cells. There are in the rec-
ords several instances of meteoric increases of free cells at other
times than at those of apparent pulses. It does not seem possible
from the data at hand to determine whether this is due to environ-
mental influences or to the accidents of collection and subsequent
handling. In the discussions of Mallomonas and Syncrypta, sugges-
tions have been made that these organisms may be stages in the
life cycle of Synura. Synura is the largest and by far the most
important synthetic organism of the winter plankton. It shares
appreciably in the winter volumetric pulses as, for example, those
of December, 1898 (Pt. I., PI. XII.).
Its fluctuations do not seem to produce any marked effect upon
the nitrates, possibly because the latter are present in excess of the
needs of Synura. In the winter of 1898 nitrates are high, 1.25
parts per million with the pulse of 1,999,500 colonies on November
29, but decline rapidly to .1 on December 13 with a fall of Synura
to 78,000. On December 20, Synura rises to 2,764,800, but the ni-
trates rise only to .35. It is evident that the nitrates are not the
only factor regulating the fluctuations of Synura.
Marsson ('00) reports Synura as abundant in the winter plankton
of lakes about Berlin, and Brunnthaler ('00) finds it in the winter
plankton of the Danube. There is, however, no recorded instance
in which Synura forms so prominent a part of the plankton of a body
of water as it does of that of the Illinois River. It may be that a
closer analysis than has yet been given the potamoplankton of other
streams will reveal its prominence there also. It is present (Kofoid
90
'95) in the summer plankton of the Great Lakes at temperatures
15 to 20 below the summer maximum of the Illinois River.
Trachelomonas acuminata Schmarda.* Average number, 1,094,-
615 ; silk, 873. This species appears in the plankton in April or May
and continues into October or November. There is no vernal pulse,
and the data are too irregular to trace the seasonal fluctuations.
The greater numbers occur during the period of maximum heat.
Excepting a single occurrence in February, this species has been
found only above 40, and its period of continuous appearance from
May to October lies above 60. It is evidently a summer plank-
tont.
Trachelomonas hispida Stein.* Average number, 1,002,115 ; silk,
1,251. This is a perennial organism, found in every month of the
year but in larger numbers during the warmer months. It was
more abundant than usual in the winter of 1897-98 following the low
water and unusual development of the previous fall. There are no
large pulses in 1898, but in 1897 there is indication of a vernal max-
imum on April 27 and an autumnal one of 85,500,000 on September
7. The data are too irregular to trace the seasonal fluctuations in
detail. There is no doubt, however, from the evidence at hand that
this is a predominantly warm-water planktont similar to the other
members of the genus.
Trachelomonas volvocina Ehrbg.* Average number, 17,672,692;
silk, 7,162. This is the most abundant species of the genus and is
found throughout the year in almost every collection. It is most
abundant from May to October, during the period of maximum
heat. There are no well-defined vernal or autumnal pulses, but
recurrent maxima during the summer are to be found in both 1897
and 1898. There are four such pulses in the former year, and in the
latter five, as follows: May 17 at 64 (14,400,000), 'june 21 at 77
(147,600,000), July 19 at 84 (86,400,000), August 9 at 83
(252,000,000), and October 4 at 71.5 (11,700,000). The periods
of greatest growth thus lie above 60 and the optimum is
near 80. None of these pulses coincides with a volumetric
maximum of the silk-net catches (Pt. I., PL XII.). They
usually follow these maxima at intervals of one or two weeks
a phenomenon often observed in other synthetic species. It may
be explained by the decrease in animals which feed upon the organ-
isms in question. These volumetric pulses are predominantly
91
animal in their composition, and when they decline the organisms
upon which the disappearing animals were feeding have an oppor-
tunity to multiply with less decimation in their ranks.
This species is one of the most abundant of the synthetic organ-
isms in the summer plankton, and next to Euglena is the foremost
among the synthetic elements of the food cycle of the plankton.
The presence of many light-colored or even colorless forms (forma
hyalina Kl.) justifies the suspicion that members of this genus, like
those of its near relative Euglena, adopt holozoic nutrition in the
presence of abundant organic matter suitable for food.
This species, as well as the others above listed, is exceedingly
variable in the proportions of the lorica, in its color, and in the
development of the neck. It is very desirable that its life history
and the full limits of its variation be determined before many more
new species are proposed in the genus.
In addition to the forms above listed, the following have been
noted as present in small numbers in the summer plankton, viz. : T.
armata Ehrbg., T. caudata Ehrbg., T. torta Stokes, T. urceolata
Stokes, and T. volvocina var. rugulosa Kl.
Uroglena americana Calkins. This species was found in small
numbers in July and September, 1897, and in January, 1899.
Uroglena radiata Calkins. This species was found in January,
1896 ; in April and May, 1897 ; and in March and April, 1898. There
was a vernal pulse of 15,279 on April 29, 1896.
Uroglena volvox Ehrbg. This species was found sparingly in the
spring plankton in 1896. Uroglena is one of the few organisms
which the usual method of plankton collection and preservation
fails to keep in fair condition for subsequent indentification.
The gelatinous matrix is easily crushed, and debris adheres to it so
as to obscure it beyond recognition. Judging from the frequency
of Uroglena in the living plankton it is very probable that the genus
is much more abundantly represented in the Illinois River than the
data at hand indicate. The genus seems to prefer the cooler waters
of autumn and spring to those of midsummer.
Volvox aureus Ehrbg. This species was found from March to
August, but in small numbers and irregularly.
Volvox globator L. This was somew T hat more abundant than the
previous species, and was found more frequently, especially during
92
1895 and 1896. It occurred from the first of May till the end of
August, but always in small numbers. It is occasionally abundant
in backwaters where there is much vegetation.
In addition to the Mastigophora above listed there were many
individuals belonging to unidentified species . They were as a rule the
smaller forms, which are not readily identified in preserved material
and under the conditions of plankton enumeration. They consti-
tute about twenty-six per cent, of the total Mastigophora enu-
merated. In silty planktons their number is relatively somewhat
larger on account of the difficulties attending the determination of
species in such material. These unidentified flagellates occur in
every collection, and are somewhat more abundant in the summer
months.
RHIZOPODA.
Average number, 55,364, including filter-paper collections;
23,826 without them. This group of Protozoa is numerically of less
importance than the ciliates or flagellates, but its quantitative
significance is greater than the numbers of individuals indicate.
This is due to the relatively large size of the Rhizopoda, and also to
the fact that plankton collections afford only an irregular and in-
complete record of the rhizopodan fauna of any body of water, and
give but an imperfect idea of the part which these organisms play
in the total economy of the lake or stream. This results from the
fact that they are as a rule largely bottom or shore-loving species,
and are generally either adventitious or temporary constituents of
the plankton.
The seasonal distribution of the total Rhizopoda in the Illinois
River gives evidence of the adventitious or temporary nature of the
contributions of the group to the plankton. There are pulses in
1898 on January 25 (66,388), February 22 (141,524), August 23
(36,800), September 27 (59,200), and November 15 (42,000), all of
which appear on rising water and are largely adventitious, their
presence in the plankton being due to the disturbances of currents,
waves, and the like. There are pulses on May 10 (49,800), June 28
(37,000), and July 19 (28,800) which cannot be traced to any
general hydrographic condition. These, as will be suggested in the
discussion of the seasonal fluctuations of individual species, are
probably due to the temporary adoption of a limnetic habit on the
93
part of some of the rhizopods,orto the appearance of limnetic forms,
varieties, or species according to the systematic value placed upon
these eulimnetic individuals. I am inclined myself to regard them
as seasonal forms of species which are predominantly of the bottom
or littoral fauna, which have multiplied rapidly under the stimulus
of abundant food. Owing to this fact, to the storage in their
tissues of the products of metabolism, such as gas and oil vacuoles
which tend to lighten their specific gravity, and to the frailer
structure of their shells under conditions of rapid multiplication,
they abandon their customary benthal or littoral habitat and assume
temporarily a limnetic distribution in the plankton where they con-
tinue to find abundant food. Their appearance here under these
circumstances is a result of their physiological condition, and with
its cessation they decline, as shown by their pulse-like occurrences.
Whatever the systematic valuation placed upon these limnetic
forms may be, there is no doubt of their occurrence. They have
appeared in every year of our operations, but were most prevalent
in 1897, a year of most stable conditions, and also in the quieter
backwaters, and on the declining spring flood or June rise when hydro-
graphic conditions are less catastrophic than those of early flood
stages. In 1897 there was a pulse of 68,400 (silk-net only) on August
8 and another of 1,268,400 on September 7, both in stable conditions
and almost exclusively of limnetic types, differing in this respect
from the pulse of 141,524 on February 22, 1898, which was pre-
dominantly of an adventitious character, resulting from the flood
of that period (Pt. I., PI. XII.). The contrast in the numbers of
Rhizopoda in the plankton during warm and cold seasons of the
year is very striking in 1897. The average per m 3 , per collection
from May 1 to October 1, that is, above 60, is 161,045, omitting all
filter-paper collections, while in the seven months of lower tempera-
tures this average is only 4,771. During the warmer period the
June rise was the only hydrographic disturbance (Pt. I., PI. XL)
to which any adventitious increase might be attributed. This con-
trast is less evident in 1898, when the summer hydrograph was more
disturbed. These larger numbers during warmer months may be
attributed in part to the greater numbers of the Rhizopoda in their
littoral habitat, and in part, doubtless, to the fact that at low water
the shore and bottom fauna are brought into more intimate relation
with the plankton, and in the river the disturbance of these regions
94
by current, waves, seines, boats, and fish make relatively larger
contributions at low-water stages to the diversification of the
plankton. In addition to these factors, however, there is abun-
dant indication that many individuals assume during the warmer
months a eulimnetic habit, and that some of the Rhizopoda
become, for the time being at least, typical, though temporary,
planktonts.
It naturally follows that in so far as the plankton is concerned,
the Rhizopoda exhibit a seasonal preference for the warmer months
above 60! Maximum numbers were attained only at the higher
temperatures save in those instances where they attend winter
floods. In a measure the seasonal distribution of the Rhizopoda in
the plankton reflects that of the group in its normal habitat ; but at
the best the picture is incomplete.
The Rhizopoda have important relations in the economy of the
plankton. They feed upon diatoms, desmids, the smaller algae, and
even the chlorophyll-bearing Mastigophora such as Trachelomonas
and Carteria. Their occurrences in the plankton do not exhibit any
striking correlation with those of the groups named. The great
pulse of September 7, 1897, for example (PI. II.), lies in a depression
of the diatoms and coincides with pulses of Chlorophycece and
Mastigophora, and that of August 10 (68,400) exhibits a similar
relation, the diatoms rising the following week as the Rhizopoda
fall. In 1898 the pulse of Rhizopoda on June 28 of 37,000 (Table I.)
culminates a fortnight after that of the diatoms and Chlorophycece
and a week after that of the Mastigophora. It thus is intercalated
between the June and July pulses of these chlorophyll-bearing
organisms (PI. II.). The Rhizopoda pulse of July 19 (28,800), on
the other hand, occurs with the coincident pulses of the three
groups named (PL II.). The immediate diluent effect of flood
waters upon the plankton combined with their tendency to increase
the number of adventitious Rhizopoda results at times in the
intercalation of their pulses with those of the chlorophyll-bearing
organisms whose relative numbers are reduced by the dilution. The
data evidently do not afford any adequate solution of the inter-
calations of the Rhizopoda with other organisms.
The Rhizopoda are very frequently found in the digestive tract
of limnetic rotifers, but I have never noted the Entomostraca feed-
ing upon them. They are important elements in the food of young
95
fish (Forbes, '80) such as the Catostomidce and some of the Silu-
rida and minnows. I have found them in great abundance in the
intestine of the adult gizzard-shad (Dorosoma), and in the contents
of the digestive tract of the German carp (Cyprinus carpio).
In the pages which follow, the seasonal distribution, or occur-
rence in the plankton, of thirty-one Rhizopoda is discussed, and
the presence in the plankton of the Illinois of twenty-eight other
rhizopodan forms which have been recognized by other writers as
of specific rank is noted. This by no means exhausts the rhizopo-
dan fauna of the environment which was the field of this investiga-
tion. A continued study of the plankton itself would doubtless
greatly extend the list of adventitious forms from the shore and
bottom, and a more careful analysis of the variants, especially in
the Difflugia globulosa-lobostoma group, would still further increase
the richness of the fauna from the systematic point of view. Hem-
pel ('99) lists sixteen species from this locality, and Penard ('02),
in discussion, remarks: "Une pareille pauvrete dans une region
riche en organismes de toute nature, est une impossibilitie mat 6-
rielle." However, neither Hempel's paper nor the present one
pretends to give a full account of all the Rhizopoda of the region.
He dealt largely with plankton collections, and the present paper
deals with them exclusively.
There is but little in plankton literature which gives with any
fulness the seasonal distribution of the Rhizopoda, or indicates that
they are of any considerable importance in the economy of the
plankton. The importance which they acquired in the plankton of
the Illinois is no doubt in part due to the nature of the environ-
ment with which we are dealing. The somewhat sporadic and
meteoric character of their appearances in our waters leads to the
inference that full seasonal analyses of the plankton of other bodies
of water at brief intervals may reveal a greater prevalence of the
Rhizopoda in the plankton than has hitherto been detected.
DISCUSSION OF SPECIES OF RHIZOPODA.
Amoeba Umax Duj. This was frequently abundant in the water-
bloom of midsummer, but was not identified in the plankton
collections.
Amoeba proteus Rosel. Average number, 342. The individuals
here assigned to A. proteus include those taken in our plank-
96
ton which belong to the type of A. radiosa Ehrbg., a type which
presents no distinctions sufficiently well-defined to separate it spe-
cifically from the first-named form. It seems probable that A.
radiosa includes small individuals of A. proteus which are not, at
the time of observation, creeping upon a substratum ; that is, they
are limnetic, floating free with filamentous pseudopodia character-
istic of that condition. Verworn ('97) has shown that A. proteus
takes the radiosa form in weakly alkaline solutions. Pond water
rich in algae may have an alkaline reaction (Knauthe, '98) in bright
sunlight. Larger individuals, distinctly referable to the A. proteus
type when taken in the plankton, possess at times the slender pseu-
dopodia of the A. radiosa type as well as the blunter ones charac-
teristic of the A. proteus form. I see no valid reason for separating
the two as distinct species. Most of the Amoeba recorded from the
plankton collections belong to the A. proteus type, the smaller ones
belonging to the radiosa type probably escaping through the
meshes of the silk net.
This species w T as found in 30 of the 180 collections examined,
being observed in all months of the year except May, November,
and December. The conditions attending its occurrence suggest
that it is not, habitually at least, an active planktont at all seasons
of its occurrence, but rather a tycholimnetic member, an invader
from the littoral or bottom fauna, or a temporary accession during
the warmer months. In the first place, both the number of occur-
rences and the numbers of individuals found are small, and the
seasonal distribution, plotted from the data of the collections of
the five years, is exceedingly irregular. Furthermore, 17 of the 30
occurrences happened on rising floods, when the fauna of the bot-
tom and shore of both the river and its tributaries is most mingled
with the plankton. Further evidence of the agency of floods in
introducing Amceba into the plankton is brought to light by a com-
parison of its occurrences in 1897 and 1898. As shown by Plates
XL and XII. , Part I., the hydrograph of 1897 is much less irregular
than that of 1898, the latter year exhibiting repeated fluctuations
in level due to floods. As a result we find Amoeba occurring rela-
tively (to the number of collections) almost twice as often in 1898
as it did in 1897. It may also be significant that Amceba was not
found in November and December, months of unusual stability in
river levels. There is, however, a suggestion in the data of distri-
97
bution (see Table I.) that Amoeba may become an active member of
the plankton during the warmer seasons, like other Rhizopoda, as a
result, perhaps, of the formation of gas or oil vacuoles in its proto-'
plasm. Of the 30 occurrences, 21 fall between ApriLlS and Octo-
ber 17, with water temperatures of 58 and 56, respectively. Of
these 21 occurrences in warm waters but 8 accompany flood inva-
sions, while all of the 9 occurrences during the colder months are in
connection with such disturbances. Finally, the maximum num-
ber per cubic meter (6,400) was found July 21 in clear waters, free
from the debris of flood invasion. In conclusion, it seems probable
that Amoeba in warmer seasons of the year (above 56) may adopt
a limnetic habit. There is, however, the possibility that local and
minor disturbances of the water due to current, waves, etc., are
the occasion of its presence in the plankton in the absence of flood
conditions. Jennings ('OOa) reports both A. proteus and A. radiosa
in the open water of Lake Erie.
The range of temperature of river water in which Amoeba was
found was from 32 to 89 the full extremes observed by us in
the river at Havana. The temperature at the maximum occur-
rence, July 21, 1897, was 82. It is perhaps significant that 14
of the 30 occurrences of Amoeba were between June 21 and Sep-
tember 6, the period of maximum heat, the river averaging
almost 80 apparently the optimum temperature for the occur-
rence of Amoeba in the plankton in this locality. The relative
numbers of individuals found in the various collections of the five
years are too irregular to suggest any conclusions as to a seasonal
cycle.
Amoeba verrucosa Ehrbg. Average number, 19. This species
was found but three times in the plankton, once each in May,
August, and September, occurring but singly, and in each case in
flood waters. It is apparently a tycholimnetic member of the plank-
ton. The temperature limits of its recorded occurrence in the
plankton were 5.8 and 82 respectively.
Arcella.
This genus is represented in the plankton by four species and
two varieties which, like most of the Rhizopoda, are exceedingly
variable, grading in some instances into each other by occasional
98
individuals which present intermediate characters. The majority
of the individuals were taken in a living condition, though many
empty shells were found. The conditions of the examination of the
plankton and the opacity of many of the shells made it impossible
to distinguish the dead shells in all cases. The records include many
dead shells.
Arcella costata Ehrbg. Average number, 48. For the purposes
of this paper I have included here all those individuals which possess
an angular or ribbed shell. Leidy ('79) refers such forms to A.
vulgaris. Individuals of this type are rare, occurring infrequently
and in small numbers. It was recorded but 18 times in the 180
collections, and the largest number per cubic meter was only 1,187.
As in the other species of the genus, the warmer months are favored,
fourteen occurrences falling in June-September in water at 70 or
above. The other four records are one each in April, October,
November, and December. The seasonal range of this form in the
plankton thus falls in the main within the period of the maximum
abundance of A. vulgaris, of which species it may be but a variant.
Arcella discoides Ehrbg. Average number, 972. This prevalent
species is not in all instances easily separated from A. vulgaris.
Indeed, even Leidy ('79) states that it graduates into A. vulgaris,
and that he views it as the variety of this species in "which the shell
presents a greater proportionate reduction in height compared with
the breadth." In the enumeration of our plankton catches, the
larger, flatter, and unornamented individuals have been referred to
this species. Both the brownish and the hyaline forms should
probably, for reasons hereafter given, be included here, and they
are so grouped in the present discussion. Thus considered, A.
discoides is the most abundant member of the river plankton be-
longing to this genus, including two thirds of all the individuals
observed.
This species occurred in almost two thirds of the collections, hav-
ing been recorded in 115 of the 180, and more frequently and in
larger numbers in the latter half of the five years than it was in the
earlier period. This is in part explained by the unusual fluctua-
tions of the river levels in 1898, during the maximum summer
occurrence of the species. Like the other species of the genus, A.
discoides has a period of maximum occurrence in the latter part of
summer, as is shown in Table I. Of the 115 occurrences, 55 were in
99
June-September, in water at or above 70, while in the remaining
eight months there were but 60 occurrences. This contrast is
heightened by the ratio of occurrences to the total number of collec-
tions, which in the period from June to September inclusive is 55
to 68 and in the remainder of the year only 60 to It2. - The num-
ber per cubic meter is also higher during this warm period, averag-
ing for a single occurrence 1,376 to 1,028 for one in the remainder of
the year. The average for the colder months falls to 850 if the
large accessions attending the floods of February and November are
omitted in the totals. The same causes efficient in determining the
summer maximum in other Rhizopoda of the plankton are doubtless
operative here, and as in A. vulgaris the impetus of the summer in-
crease is carried over into the autumn, causing a slight increase in
numbers as compared with the numbers at corresponding temper-
atures in the spring months. It seems probable that high temper-
atures favor its occurrence in the plankton, not, however, directly,
but because of greater abundance of food under those conditions,
greater metabolism, and the storage of the products as oil or gas
vacuoles which tend to lower the specific gravity and thus to bring
the animal into the plankton.
The adventitious occurrence of A. discoides in the plankton is
shown by the fact that 45 of the 115 occurrences are with rising
flood waters. The greater part of them lie in the colder months;
in fact, nine tenths of the occurrences between October and May are
correlated with flood movements. For reasons above given, how-
ever, A. discoides may be regarded as temporarily adopting a lim-
netic habit during warm months as a result of its physiological
condition ; at least many individuals of the species exhibit this habit
during the warmer months. The data do not indicate that the
open water is at any time the center of distribution of the species.
There are no indications of recurrent pulses in the species and,
as might be expected in case of adventitious planktonts, but little
evidence of a characteristic seasonal distribution. There is some
evidence that the summer is the period of most active multiplica-
tion, and that an exceedingly transparent and hyaline form other-
wise resembling A. discoides is the young of this species. In 1898 .
separate records were kept of the two types with the result that they
were about equally abundant 24,159 and 26,387 for the brown
and hyaline types respectively.
(8)
100
With but few exceptions the seasonal distribution exhibited by
the hyaline form was very similar in time and numbers to that of the
brown form. Both occurred more frequently and in larger numbers
in the warmer months, and irregularly and in small numbers in the
colder waters. Both entered in larger numbers with flood waters.
The differences though slight are suggestive. The hyaline form was
less frequent than the brown both in occurrences and numbers dur-
ing cold weather, and summer floods sometimes brought a rela-
tively larger number of the hyaline type. These are conditions
that might be expected if the latter is only the young (that is, the
daughter organism occupying the new shell after fission of the oc-
cupant of the old) of Arcella discoides. In warmer months food is
more abundant and, presumably, fission more frequent. For this
reason the young individuals abound at that time. Owing to the
difference in the specific gravity of the two, the hyaline type is
more readily transported by flood waters. Though not con-
clusive, the data here presented seem to favor the view that the
hyaline form is only a stage in the life history of the individual
Arcella discoides.
The species A. artocrea Leidy and A. polypora Penard occur also
in our waters, but were included with A. discoides in the enumera-
tion. Typical representatives of these species are not, however,
present in any numbers
Arcella mitrata Leidy was found but once on Aug. 1, 1895, in
small numbers, at 78.5.
Arcella stellata Perty. Under this designation are included only
those individuals which have well-defined prolongations on the
margin of the shell. Only a single occurrence in small numbers
(48 per cubic meter) was recorded for the typical A . stellata July
29, 1895, at a temperature of 75.5.
Arcella vulgaris Ehrbg. Average number, 1,098. This species
is somewhat more abundant than A . discoides, but occurred in fewer
collections. It is a somewhat common planktont, whose seasonal
distribution exhibits some irregularities attributable in part, as in
the case of other members of the genus, to flood conditions. It was
found in 61 of the 180 collections examined, and in approximately
one third of those made in each year, excepting in 1894, when it
was not recorded, and in 1898, in which year it was found in about
101
half the collections, the river levels for this latter year being subject
to more than the usual disturbance.
Arcella vulgaris is found throughout the whole year, with a
marked predominance of occurrences during the warmer months,
June to September inclusive, for during this period, irr which a total
of 68 collections were made, this species was found in the plankton
34 times. If the month of October be included, the ratio is 44 oc-
currences in 83 collections, while in the remaining 97 collections,
from November to June, only 17 occurrences were recorded. Of
the 10 occurrences in October, 7 were in water at or above 55. The
season of frequency in the plankton thus ranges from June
through October. In both frequency of occurrence and in numbers
of individuals (see Table I.) there is an apparent maximum in
August, preceded by an increase in June and July and followed by
a decline in September and October. Arcella vulgaris thus seems
to be a late summer planktont. The continuance into October
may in part be due to the temperature conditions above cited, and
perhaps also to constant seining of the river by fishermen in the
low- water stages at that time, causing repeated disturbances of the
bottom and shores, where Arcella habitually lives. This maximum
frequency of Arcella during the warmer months in the plankton
is, however, probably due to the formation of gas or oil vacuoles in
the plasma under the conditions of higher temperatures. Their
flotation is thus facilitated, and they become, in a way, semi-active
but temporary planktonts.
That floods are also in part responsible for the presence of Arcella
in the plankton is evident from the fact that 32 of the 61 occurrences
come with rapidly rising waters, or shortly after rapid rises, during
the interval of rapid decline. The larger numbers of individuals
also appear in flood- waters, occurrences of more than 1000 per cubic
meter happening 10 times with floods to only 4 in more stable
conditions. The maximum occurrence, 25,272 per cubic meter,
came with the flood of February, 1898, indicating the presence of
this species in large numbers, even under winter conditions, in some
local environment tributary to the flood plankton.
The average number per cubic meter in the 61 collections con-
taining Arcella was 1,260; and the maximum, 25, 272, as above noted.
This species occurred in only 10 collections in stable conditions of
the river, when the temperature of the water was below 55. The
102
average number of individuals in these cases was, however, only
230 per cubic meter as against 1,443 when the temperature was
above 55, or, if below, when floods prevailed. The seasonal and
numerical distribution of occurrences and individuals alike point
to the agency of floods and higher temperatures in the introduction
of Arcella into the plankton from its usual habitat, the bottom and
the shore.
This species occurred in water ranging in temperature from 32
to 89. Being a bottom form, the plankton data do not afford a
satisfactory basis for determining its true seasonal distribution and
optimum temperature. The maximum number found, 25,272, was
in water at 32; but this was an isolated occurrence in a flood, and
serves only to illustrate the irregularity of distribution in the
plankton of tycholimnetic organisms.
Centropyxis aculeaca Stein. Average number, 570. This species
has appeared in collections in every month of the year, but its
sequence is frequently interrupted and its numbers are quite irregu-
lar. Practically without exception all the larger occurrences attend
rising flood waters. It is evidently adventitious at all seasons of
the year.
Centropyxis aculeata var. ecornis (Ehrbg.) Leidy. Average
number, 604. In former years this species was less frequent than
the preceding species. Its appearances in the plankton tend to
coincide with those of C. aculeata (Table I.), and are doubtless due
to the same causes. Thus in the February flood of 1898 there is a
pulse of 12,636 of C. aculeata and one of 9,477 of var. ecornis.
C. l&vigata Penard seems to be identical with this variety. The
data concerning both C. aculeata and its variety ecornis are too
irregular to throw any light on the seasonal cycle of these adventi-
tious planktonts.
Cochliopodium bi limbo sum (Auerbach) Leidy. Average number,
1,384. This species was found in the plankton during 1898 in
irregular numbers in 27 of the 52 collections. The distribution of
the occurrences affoids indubitable proof of their close dependence
upon flood waters. In 15 of the 27 cases Cochliopodium appeared
with a rising river, and in all but 6 cases, in periods of considerable
movement in river levels (cf. Table I. with PL XII., Pt. I.), such
as the rising flood of January and February and the repeated minor
103
fluctuations of August and the following months. The year 1898
was one of unusual irregularity in the hydrograph (Pt. I., PL XII.),
especially at the lower stages of the river, at which times this
rhizopod appeared most frequently. Its maximum occurrence,
20,898 per cubic meter on Jan. 25, accompanied a rise" of 0.6 of a
foot in 24 hours. At other times the numbers range from 100 to
8,000 per cubic meter, their irregularity affording additional ground
for regarding this species as an adventitious planktont.
Cochlio podium was present in water ranging from 32.1 to 89,
the maximum number observed being found in water almost at the
freezing point, when the river was full of running ice. That this
is the optimum temperature for this organism is not, however, to
be inferred, since, as has been shown above, this species is adventi-
tious in the plankton. Plankton collections do not afford adequate
data for determining the seasonal cycle of the organisms habitually
living upon the bottom. This species was not found, though careful
search was made for it, in the winter collections of 1899. Its
absence from the records of years previous to 1898 may in part be
due to a failure to observe it in the silt-polluted collections in which
it is most apt to occur.
Cyphoderia margaritacea Ehrbg. Average number, 198. This
species has occurred in every month but February. In 1898, the
majority of the occurrences and three fourths of the numbers ap-
peared between May 1 and October 1 at temperatures above 60. It
was never abundant at any time, though there is this indication of
its increased numbers during the warmer season. It is not an im-
portant element in our plankton. Apstein ('96) found it somewhat
irregularly in the plankton of German lakes. In our waters it
exhibits no marked dependency upon floods for its presence in the
plankton, though it is probably capable of assuming the limnetic
habit in the warmer season.
Cyphoderia trochus Penard appeared occasionally with the pre-
ceding form, from which it is distinguished by its conical horn on
the fundus and by its larger scales.
Difflugia.
This genus is the most abundant one of the Rhizopoda in the
plankton of the Illinois River, and is a factor of quantitative
104
importance in its economy. It includes a number of forms notorious
for their variability and for the difficulty with which specific dis-
tinctions can be applied. I shall discuss the species as they were
enumerated, and shall correlate my work with Penard's ('02) recent
elaborate analysis of the species so far as I can with the aid of my
notes in the absence of the collections. Opinion as to the validity
of the species is expressly withheld excepting in those instances in
which it is formally stated.
Difflugia acuminata Ehrbg. Average number, 315. This spe-
cies has occurred in every month of the year and in 83 out of 180
collections. In 1898, two thirds of the occurrences and three fourths
of the individuals were taken between May 1 and October 30, at
temperatures above 70. In this year there are six recurrent pulses
from June to November, but all but one of these are found on rapidly
rising flood waters, and they bear no constant relation to the pulses
of diatoms previously noted, with which in some instances they
are intercalated, though this is not regular or constant. Similar
tendencies to appear with floods and in greater numbers and more
frequently in summer can be detected in records of other years. It
was more than twice as abundant in 1896 a year of interrupted
hydrograph (Pt. I., PI. X.) as in 1898. This is one of the larger
and heavier rhizopods, and its occurrence in the plankton is doubt-
less adventitious, due to floods and currents, and its greater numbers
and frequency in the summer may result from its greater abundance
at that season in its natural habitat, the shore and bottom, and
perhaps, also, from its lighter specific gravity during the warmer
season. An illustration of this appears on the rising flood of June,
1897, when the maximum number recorded (10,000 per m. 3 ) oc-
curred.
The shell of this species is exceedingly variable in size, constitu-
ent particles, and proportions. A number of forms separated by
Penard ('02) and others as distinct species were grouped under D.
acuminata in the enumeration. The greater number of these belong
to the type designated by this name by Penard ('02). D. acuminata
var. inflata Penard and the somewhat similar D. elegans Penard are
not uncommon. D. acuminata var. umbilicata Penard, D. elegans
var. teres Penard, D. curmcaulis Penard, D. lance olata Penard, and
D. scalpellum Penard occur also, but are rare.
105
Difflugia bicuspidata Rhumbler. Average number, 76. A sep-
arate record was kept of this bicuspid type in the later years of
our collections. Penard ('02) regards it as a synonym of his D.
elegans, though it would seem to be as worthy of specific distinction
as many other variants to which he accords this raak_ It varies
greatly in the relative development of the accessory "horn,"
which is sometimes but a mere elevation near the base of the main
horn. Individuals with equal and symmetrical horns represent the
other extreme. In a few cases tricuspid individuals have been seen,
evidencing a tendency to vary towards the type found in D. varians
Penard and D. fragosa Hempel.
This form was about one fourth as abundant as D. acuminata,
and eight of the ten occurrences fall between May and October, usu-
ally with D. acuminata and presumably for the same reasons.
Difflugiq constricta Ehrbg. Average number, 46. This species
occurs irregularly at all seasons of the year without marked prefer-
ence for the warmer months, and often, but not always, with flood
waters. It occurs throughout the whole range of temperatures, and
the largest number (2,778 per m. 3 ) appeared during the decline of the
spring flood. Data are too infrequent to establish any seasonal
routine.
This species varies greatly, and is connected by an unbroken
series of variants with the genus Centropyxis. Penard ('02) also
notes the existence of this connection, and states that after careful
search he was unable to find any constant distinction w r hich would
suffice for its separation. In my enumeration only the elongated
and smooth individuals were referred to this species. The spinose
forms were referred to Centropyxis aculeata, and tho.se similar in
form to the spinose type ; but those free from spines, to C. aculeata
var. ecornis.
Difflugia corona Wallich. Average number, 36. In 1896, when
the hydrograph was much disturbed, the average number was more
than twice as great. This superb species was found in every month
of the year except December, but never in large numbers. Its
large size (200-300 fi), and its heavy shell militate against its pres-
ence in the plankton, and its occurrences are irregular and its num-
bers few. There is no marked preference for warmer months, and
four fifths of its occurrences are in rising flood waters. It is plainly
106
an adventitious planktont. The data are too irregular to trace its
seasonal distribution.
As a species it is as well defined as any in the genus. It is not in
our waters connected by intermediate forms with other species. Its
assignment to D. lobostoma by Schewiakoff ('93) is not in my opinion
justifiable unless we regard all forms of Difflugia as belonging to one
species.
Difflugia fragosa Hempel. Average number, 25; in 1896 over
100. This species occurred in every month of the year but Febru-
ary, though three fifths of the records and the majority of the in-
dividuals were found between May and October at temperatures
above 60. The data are too irregular to trace the seasonal history
of the organism, but they suffice to suggest the agency of floods at
all times and of high temperatures during the summer, as factors
in the occurrence of the species in the plankton. The shell of this
form is relatively to that of other species rather heavy, and this fact
combined with the irregularity of its occurrence seems to justify
the conclusion that it is largely adventitious at all seasons of the
year.
The species exhibits a great deal of variation in the development
of the central spine Hempel ('99, Fig. 1) and in the number and
arrangement of spines in the accessory circlet. The mammillate
form of the central spine figured by Hempel is not usually present.
Individuals in which the central spine is but feebly developed seem
to connect this species with D. varians, recently described by
Penard ('02). Otherwise, and in our waters, the species is well
delimited.
Difflugia globulosa Duj. Average number. 7,194; in 1897,
47,329, the larger number in this year being in part due to a remark-
able pulse of 1,240,000 early in September. This is the most
abundant of all the rhizopods in our plankton, occurring most
frequently and in largest numbers. It is found in every month of
the year, and in 1898 appeared in every collection except four in De-
cember. With a few exceptions in the autumn of 1898 (Table I.),
no large development (exceeding 10,000 per m. 3 ) has taken place
earlier than May or later than September that is, at temperatures
below 60. The occurrences are most continuous and the numbers
of individuals are largest during the warmer period between the
months named. The largest pulse, that of 1,240,000 on September
107
7, 1897, was at 80. A pulse of 48,000 on November 22 at 40 gives
evidence of considerable range in adaptation to temperatures.
In Table I. the seasonal distribution of D. globulosa is given in
full. It differs from that of previous years mainly in the fact that
the summer pulses do not here have the amplitude reached in other
years; for example, in 1896 (252,000) and 1897 (1,240,000). It is
characterized by considerable irregularity caused by somewhat
abrupt pulses at irregular intervals. A comparison of these occur-
rences with the hydrographic conditions (Pt. I., PL XII.) indicates
that in the colder months increase in numbers in the plankton at-
tends flood waters only, as, for example, in January, February, late
October, and November. In the summer, pulses may also come
with floods. For example, that of 252,000 on May 25, 1896, ap-
peared on the upward slope of the June rise of the year, and that
of 80,000 on June 28, 1897, came with the belated June rise of that
year. On the other hand, some of the minor fluctuations appear
on declining floods, and the maximum one of our records, that of
Sept. 7, 1897, came in the midst of the most prolonged period of
stable low water (Pt. I., PL XI.) found in the six years of our
operations. From these facts it is evident that floods are efficient
in increasing the number of D. globulosa in the plankton, and that
the amplitude of the pulses to which they contribute is much greater
in the warmer months (above 60) than in the colder ones as a
result, perhaps, of the greater numbers present in their normal
habitat, the shores and bottom, and also as a result of their readier
flotation at this season. In so far as their presence is due to floods
they are adventitious. On the other hand, it is very probable that
they become temporarily eulimnetic in habit during the summer
months. The evidence for this lies in their greater numbers in a
period which is predominantly one of greater stability. Thus in
1898, in the 22 collections between May 1 and October 1, the average
number present is 9,731, while in the remaining seven months of
colder weather the number is only 5 ,200. Additional evidence arises
from the fact that pulses of unusual magnitude have occurred quite
independently of any factor such as flood or other disturbance which
might cause their adventitious introduction into the plankton.
Thus on Sept. 7, 1897, there is a symmetrical pulse whose rise and
decline occupy four weeks, as shown in the following table. The
total change in river levels in this period of four weeks (Pt. I., PL
108
Date
Number per m. 3
Turbidity
(in meters)
Silt
(in cm. 3 )
Stage of
river above
low water
August 24
4,800
.37
.15
1.8
August 31
112,000
.33
.19
1.8
September 7
1 240,000
.15
.45
1.8
September 14
106,000
.33
1.04
2.0
September 21
800
.35
trace
2.0
XL) was only a fall of .1 and a rise of .2 of a foot changes due to
wind and the operation of the locks in the dams at either end of the
pool. The estimated percentage of silt is near the minimum from
a trace to 5 per cent. and the turbidity was no greater than is
customary (Pt. I., Table III.) in our waters during periods of abun-
dant plankton such as this (Pt. I., PI. XI.). Beyond the presence
of these rhizopods there was nothing in the plankton to suggest
that the bottom had been stirred up any more than usual. No
environmental factor is apparent to which we can attribute this
wave of Difflugia in the plankton. It is due, I believe, to their own
physiological condition. This was a time of prolonged low water
and great sewage contamination, and of remarkable development of
water-bloom, chlorophyll-bearing flagellates, unicellular algae, and
some diatoms, all elements in the food of Difflugia. In the open
water Difflugia could find abundant sustenance and thus maintain
itself there. It is not strange, then, that we find it in these warm
waters, richly charged with its food, assuming for the time a eulim-
netic habit, perhaps as a result of rapid growth and lighter shells, and
of increased metabolism with reserve products which lighten the
specific gravity and so facilitate flotation.
This species is found throughout the whole range of temperatures,.
There are indications that its optimum lies above 60, and perhaps
near the maximum, 80. This may, however, be the result of the
effect of temperature upon the food supply of the organism. In any
case the plankton data can not suffice to follow the complete seasonal
cycle of an organism which is either an adventitious or but a tem-
porary constituent.
109
The question of specific limits and variation in this organism
is one of exceeding difficulty, and I see no satisfactory solution for
it until some one attacks the problem by a study of the variation
by modern quantitative methods, and endeavors by breeding under
control to establish the limits of variation within the^iormal range
of seasonal changes of the environment. When this is done, some
more satisfactory criterion for species in this group of planktonts
will be feasible than the present condition affords, in which slight
differences from previous descriptions are held to be valid for specific
distinctions. Thus, in recent years, species of plankton Difflugia have
been described by Heuscher ('85) (D. urceolata var. helvetica) from
Swiss lakes ; by Zacharias ('97) (D. hydrostatica) from Lake Plon ; by
Garbini ('98) (D. cydotellind) from Italian lakes; by Levander ('00)
(D. lobostoma var. limneticd) from Finnish waters; and by Min-
kiewitsch ('98) (D. planktonicd) from Russian waters. All of these
forms occur in the Illinois River, and there are others equally worthy
of specific designation in our plankton as yet undescribed. They
occur most abundantly at the times of the pulses, especially of those
in stable conditions. In my opinion they are all mere limnetic
varieties of D. globulosa or D. lobostoma, the form of the shell and its
constituent particles being modified by the habit of life in which
these individuals of the seasonal cycle are found. They occur at
times of abundant food, rapid multiplication, and limnetic environ-
ment. Their shells are accordingly lighter, more chitinous and
transparent, and the foreign particles adherent to them partake of
the nature of those of the silt in suspension. This, however, is
merely an opinion based upon an examination of the statistics of
occurrences, and upon the work of plankton enumeration in which
all individuals must be assigned to some species. This is at least a
different point of view from that of the systematist, who may, per-
haps, lay more stress upon divergences from described types and
less upon links connecting such variants. For the sake of genuine
progress in the science it would seem to the writer extremely desir-
able that more attention be given to the question of variation and
less to the description of new species under criteria now in vogue. It
may be desirable, indeed necessary, to distinguish such forms in the
plankton. It would be both safe and conservative to designate
them as forms, or, at the most, as varieties.
110
The location of the pulses of D. globulosa bears no constant rela-
tion to those of other organisms, owing, in part, at least, to the
irregularities of the floods upon which some of them seem to depend.
The great pulse of Sept. 7, 1897, is intercalated between two pulses
of diatoms and other chlorophyll-bearing organisms, and some
others bear a similar relation to their food supply, while some co-
incide with an increase in these synthetic organisms (cf. Table I.
and PL II.).
Difflugia globulosa and the following species were reported by
Smith ('94) in the plankton of Lake St. Clair; by Jennings ('OOa) in
that of Lake Erie; and were common in the plankton of Lake
Michigan (Kofoid '95). Difflugia of the forms included here under
D. globulosa and D. lobostoma have been reported by many authors
from various European lakes and rivers, but in no reported instance
do they reach the numbers or importance in the plankton that they
do in the Illinois. Full records of their seasonal distribution may,
however, bring such importance to light.
Difflugia lobostoma Leidy. Average number, 1,158. In the
total of all collections it is about one fifth as abundant as D. globu-
losa. Like that species it occurs throughout the whole year in
almost every collection (Table I.), and the fluctuations in its occur-
rence follow very closely those just described for D. globulosa in the
direction of their movement. The amplitude of the pulses is less, as
a rule, and their culminations and limits are coincident, or at least
approximate. Thus, on Sept. 7, 1897, D. lobostoma attains only
24,000, and the pulse of D. globulosa on June 28 (80,000) is attended
by one of 96,000 in D. lobostoma in the next collection, on July 14.
There are in this species also the same influx into the plankton
with floods, and increase in numbers at temperatures above 60.
There are 954 per collection per cubic meter below this temperature
to 1,436 during the warmer months in 1898. There are also pulses
during the warmer months, in stable conditions, coincident with
those of D. globulosa. Similar causes presumably contribute to
these results in both species.
Difflugia lobostoma is also exceedingly variable in proportions, in
the texture of the shell and the degree of incision, and in the num-
ber of lobes about the mouth. Two, three, and even four have been
noted, and they vary greatly in depth, in regularity, in perfection
of their development, and in the structural border which sometimes
Ill
forms their margin. Chitinous, brownish, or more or less trans-
parent shells are abundant when pulses occur. Forms which
connect this species with D. globulosa have been observed. In-
cluded with D. lobostoma are forms which have since been described
by Penard ('02) as D. gramen, D. gramen var. acETom, and D.
lithopUtes, though I have not found in the Illinois plankton any of
the last-named with the peculiar tipped horns found by Penard
upon many individuals of his species.
Difflugia pristis Penard (?). A small Difflugia was found occa-
sionally in the filter-paper collections in the colder months, but
only from November to March. It was often dark, or even blackish,
resembling in this respect Penard's D. pristis. Individuals not
thus darkened approach more nearly D. fallax Penard and D. puleoc
Penard.
Difflugia pyriformis Perty. Average number, 368. This species
occurred in every month except January, but generally in small
numbers and irregularly. The largest number taken 12,000, on
May 25, 1896 came with the flood at that time (Pt. I., PI. X.), and
all the large occurrences of 1898 came with rapidly rising water
(cf. Table I: and Pt. I., PI. XII.). There are no indications of pulses
during stable conditions, and we must conclude that the species is
purely adventitious in our plankton. It is one of the largest species
with a heavy shell, and its flotation is impeded thereby.
This species is exceedingly variable. The following varieties or
variants, given specific rank by some writers, have been noted, and
are included with D. pyriformis in the enumeration: D. pyriformis
var. nodosa Leidy, D. pyriformis var. daviformis Penard, D. pyriformis
var. venusta Penard, and D. pyriformis var. lacustris Penard. A
more slender and smoothly contoured form than the last is not
uncommon.
D. capreolata Penard and D. bacillifera Penard were also found ,
but are rare.
Difflugia rubescens Penard was taken but once on May 25, 1896.
Difflugia tuberculosa Hempel was also found but once in the
planktons enumerated, though Hempel ('99) reports it as appearing
occasionally from August to November in 1895.
Difflugia urceolata Carter was taken only in April and May, 1896,
in small numbers at temperatures of 66- 80.
112
Dinamcsba mirabilis Leidy was found in the plankton but
once Apr. 12, 1898, in small numbers, at 52.
Euglypha alveolata Duj. was found in small numbers in the
plankton, but only on Nov. 1, 1898, and March 14, 1899, at tempera-
tures of 45 and 36.
Euglypha ciliata Ehrbg. appeared in the filter-paper collections
in 1897, in July, August, and November, in small numbers at tem-
peratures ranging from 80 to 48. This is said by Penard ('02) to
be predominantly a sphagnum species, but widely distributed
elsewhere in small numbers.
Euglypha Icevis Perty. This minute rhizopod was found in the
filter-paper collection of Oct. 4, 1898, at 72.
Nebela collaris Leidy was found only once on June 25, 1898, at
32.
Pontigulasia incisa Rhumbler. This curious rhizopod occurred
in the plankton in July and August, 1895, and again in August and
September, 1897, at temperatures of 75- 85. Both occurrences
were in stable conditions, and the temporary adoption of the lim-
netic habit is suggested by their appearance at these times. Two
other records in 1897 on March 22 and November 9, at 44 and
50 extend the seasonal range of the species. These occurrences
attended rising water and were apparently adventitious.
Trinema enchelys (Ehrbg.) Leidy. Average number, 158. This
little cosmopolite rhizopod of the sphagnum fauna was found but
eight times in the plankton. The individuals observed were all dark-
ened by the granular food vacuoles to such a degree that structural
details were obscured. It was noted only in the somewhat turbu-
lent years of 1898 and 1899, though on account of its small size and
the obscurity of its structure it may have been overlooked in previ-
ous collections. The few occurrences are insufficient to establish
any seasonal routine. They were at both extremes of the tempera-
ture range and in all seasons but spring, with a predominance i 'n late
summer and fall. The species is evidently adventitious in the
plankton, as shown by irregular distribution and small numbers, and
by the fact that its occurrences coincide in all instances but one with
rising water.
113
HELIOZOA.
The Heliozoa of the plankton of the Illinois are few both in
number of species and of indiyiduals. They apparently play but a
small part in the economy of the plankton. The average number
for 1898 was but 4,883. Their occurrences are confined -in ^the main
to midsummer and early autumn. But four species were identified,
though several others remain undetermined for lack of sufficient
material, especially of the living forms. Apstein ('96) reports
Heliozoa in considerable numbers in German lakes, with maxima
in July- August. It is probable that these delicate forms are fre-
quently crushed in manipulation or hidden in silt in our collections.
DISCUSSION OF SPECIES OF HELIOZOA.
-Actinophrys sol Ehrbg. Average number, 62. This species
occurred irregularly from April to the early part of November at
temperatures above 46. It was recorded most frequently in the
latter part of the summer, the largest number (28,000) appearing
Sept. 7, 1897, at 80.
Actinosph&rium eichhornii (Ehrbg.) Stein. Recorded a few times,
from July to October, at maximum temperatures (75- 80), but
always in small numbers.
Endophrys rotatoriorum Przesm. This heliozoan (?) has been
recently described by Przesmycki ( '01) as parasitic, during a part of
its existence, in Philodina and Hydatina. A parasite resembling
this parasitic stage of Endophrys was observed by me in a bdelloid
rotifer (Rotifer tardus) on several occasions, but it was never abun-
dant, nor was its connection with any free-swimming condition
noted. The heliozoan affinities of this organism seem very ques-
tionable.
Nuclearia delicatula Cienk. Average number, 4,760. This
species in 1898 appeared first on June 21, attained a pulse of 78,400
on August 9 at 82 and another abrupt one of 65,600 on September
27 at 73, and made its last appearance October 25 at 48. Occur-
rences in previous years are confined to midsummer. Its optimum
conditions of temperature obviously lie near the summer maximum,
and its lower limits near 50. Its appearance in the plankton is
not traceable to flood conditions, and it is apparently eulimnetic
in our waters.
114
Hempel ( '99) reports Raphidiophrys pallida Ehrbg. and R. elegans
Hertwig and Less, in the plankton of Quiver Lake adjoining the
river, and I have found an undetermined species of Acanthocystis
and a small heliozoan resembling Nuclearia in the river plankton.
SPOROZOA.
Triactinomyxon sp. In the plankton collections of each year
there have been found free limnetic spores which unquestionably
belong to that highly aberrant and peculiar group of organisms
described by Stole ('99) as Actinomyxidia and regarded by him as
Mesozoa, but later referred by Mrazek ('00) Caullery and Mesnil
('04), and Leger ('04) to the Myxosporidia. The organisms de-
scribed by Stole were parasitic in fresh-w r ater oligochaetes, and it is
not improbable that the limnetic spores taken in our plankton
collections are derived from parasites in some of the numerous
aquatic oligochaetes, or other invertebrates, found along the bottom
and shores of the stream.
The species here referred to Triactinomyxon differs in some
details from T. ignotum Stole. It was found in the course of the
six years at least once in every month of the year, but most regularly
in May-September, and rarely and in small numbers in the colder
months. Its transparency and long, slender, radiating, tripod-like
arms give it a typically limnetic habit.
Actinomyxidia, gen. et sp. indet. Clusters of eight, or less,
cylindrical spores radiating from a common center and bearing a
marked resemblance in structural features to those of Triactinomyx-
on, but lacking any anchor-like projections, were found sparingly
in the plankton in June-September.
The distinctively limnetic habit of these spore stages in the life-
history of these parasites is unique among the Sporozoa, and has
not, to my knowledge, been before noted.
Many of the rotifers of the summer plankton, especially Brachi-
onus and an occasional Asplanchna, have been heavily parasitized
internally by small sac-like bodies, often pear-shaped, with the
smaller end attached to the lorica, or of spherical or flattened form.
They occur in such numbers at times as to be a menace to the
rotifer population. They are usually most abundant in any given
species at the time of, or subsequent to, its maximum occurrence. It
115
was not unusual to find as high as ten or fifteen per cent, of the
individuals parasitized, and a number of empty loricas bearing addi-
tional testimony to their destructive agency.
Bertram ('92) describes these structures as " parasitische
Schlauche" in the body cavity of rotifers, and Przesmycki ('01)
works out their life history, and describes the organisms as Dimoe-
rium hyalinum, but does not designate their systematic position or
affinities. There are, however, marked suggestions of sporozoan
affinities in the organism found in the rotifers of the Illinois plankton,
which seems to be identical with that described by Przesmycki ('01).
Obviously it is difficult to take a census of such internal para-
sites. A record was kept, however, of the number of parasitized
individuals in each species of rotifer, and references will be made to
these results in the discussion of the hosts. Dimcerium appeared
in both summer and winter rotifers, and its seasonal distribution
naturally depends upon the number of available hosts. It was in
consequence most abundant during the midsummer and autumn
months.
CILIATA.
Average number, 15,812,346, including filter-paper collections.
If these be excluded and the silk catches only averaged, the number
will fall to less than a tenth of this sum. The ciliates are found in
the plankton of the Illinois throughout the whole year, and as' a
whole they do not exhibit any common seasonal predominance. The
analysis of the distribution of the individual species which follows,
exhibits two diverse tendencies which affect the distribution of the
totals. These are the vernal and autumnal pulses of the Tintinnidce,
represented by Codonella cratera and Tintinnidium fluviatile, and
the autumnal-winter occurrence of a large number of species during
the height of the sewage contamination and bacterial development.
The dominant species in this ciliate wave are Carchesium lachmanni,
Epistylis, Amphileptus, Lionotus, Plagiopyla nasuta, Glaucoma
scintillans, Stentor niger, and 5. c&ruleus. Some species, as Halteria
grandinetta, have a wider seasonal distribution, and others, as
Vorticella, Trichodina, Zoothamnium, Pyxicola affinis, and many
others, are adventitious in the plankton. Still others, as Rhabdo-
styla, Cothurniopsis vaga, Operculana, and similar peritrichan
parasites, are passive members of the plankton. The actively
(9)
116
limnetic ciliates are very few. As such we may include Codonella
cratera, Tintinnidium fluviatile, and possibly Stentor niger. Car-
chesium lachmanni and Epistylis enter the plankton only in the
form of detached and often moribund zooids, and thus are not
typical planktonts, though of quantitative importance in our plank-
ton in the colder months. A large number of species not here
reported occur in our collections made elsewhere than in the
river channel, especially in places where the decay of large quan-
tities of organic matter is in progress. This is not a condition
normally found in the open water of lakes, though it may occur
along their shores, where vegetation is found, or in regions of
sewage contamination. In the waters of the Illinois, on the
other hand, the current, combined with sewage and industrial
wastes and the organic detritus from the richest of fertile prairies,
provides a suitable environment, even in the open water, for
the support of a ciliate fauna of a magnitude somewhat unusual
in fresh- water plankton. This fauna is present also in the back-
waters, but is less abundant there than in the river itself. These
species occur in greatest numbers of individuals in our plankton dur-
ing the winter months at minimum temperatures, rising in November
as the temperature falls below 50, and declining again as it rises to
this point in April. As shown by the bacteriological investigations
of Jordan ('00) and Burrill ('02 and '04), the bacterial pulse attend-
ing the decay of the sewage and wastes at Peoria does not reach
Havana during the warmer months (see table on p. 231, Pt. I.),
but when temperatures pass below 50 in November the increase in
bacteria is marked. The decay is less rapid at low temperatures,
and the process is still going on when the water in the channel
passes Havana during the prevalence of low temperatures, and the
ciliates that thrive in such an environment abound in the plankton
at that time.
The temperature limits of these ciliates of the period of bacterial
development thus seem to lie between 50 and 32. An examination
of the plankton in the river at several points between Peoria and
Havana at intervals throughout a year, will reveal how far the
component species of this ciliate fauna are governed in their seasonal
distribution in the plankton at Havana, respectively, by conditions
of temperature and by the state of sewage contamination. The
work of Roux ('01) upon the Ciliata about Geneva would seem to
117
indicate that many species of the fauna of stagnant water are more
abundant in that region during the winter months. Owing to the
difference in food conditions attendant upon the increase of sewage
and bacteria during the colder months in the Illinois River, it is
impossible to determine from the data at hand the relative efficiency
of the two elements of temperature and food in regulating the
seasonal occurrences of our ciliates.
Here, as elsewhere, the disastrous effect of sudden floods can be
traced. The number of ciliates (Table I.) drops as floods rise, and
recovers as the waters fall again. For this reason the winter occur-
rences of the total ciliates are subject to considerable disturbances
in the winter floods of the several years. The combination of the
two methods of collection and of the two groups of ciliates, typical
and adventitious, causes further irregularities (Table I.) in the sea-
sonal distribution of totals.
In the Illinois River, for reasons given above, the Ciliata occupy a
place in the economy of the plankton of more than the usual im-
portance. They feed principally upon bacteria, decaying organic
matter, and the smaller algae, and are themselves eaten by the
rotifers. I have found no evidence that they are utilized by the
Entomostraca. They thus become active agents in the reduction
of sewage and in the destruction of the bacteria of decay, in the
purification of sewage-laden waters, and in the transfer of the matter
in sewage to higher forms of animal life.
The ciliates found in the Illinois include all the important species
reported in the plankton of fresh water, and the list is somewhat
larger than hitherto recorded in quantitative plankton collections
in river or lake waters. These organisms escape readily through the
silk net by reason of their small size, and in some instances the
larger species, by reason of their mobility and flexibility, escape
through the silk where less motile organisms of equal size are re-
tained. By experiment I have found that well -shrunken silk
bolting - cloth whose meshes average about 30-45/1 will not retain
Paramecium whose diameter is 40-70/*. It may be that supple-
mentary methods of collection which will correct the error of leakage
will show that the Ciliata are of wider occurrence in the plankton
than has hitherto been found to be the case.
118
DISCUSSION OF SPECIES OF CILIATA.
Amphileptus spp. Average number, 630. Amphileptus is a well-
defined winter planktont in the river at Havana, and it affords a
striking instance of the interdependency of organisms in the plank-
ton. It feeds upon the heads of Carchesium lachmanni, engulfing the
head in situ and encysting during digestion. Such heads, joined to
the colony or free in the plankton, have been found in our waters. Its
seasonal distribution at Havana is almost identical (Table I.) with
that of Carchesium, upon which it feeds. Thus in 1897-98 Car-
chesium was continuously present in the plankton from October 26
to May 10, with a pulse on December 7 of 283,800, and one on
February 8 of 197,600. Amphileptus appears October 26 ; continues,
with interruptions, to May 17; and has pulses December 7 and
January 25, the latter reaching 13,545. In 1898-99 both appear
early in October and have coincident pulses on November 22 and
January 24. In 1895-96 the interdependence is even more striking,
Carchesium reaching a greater development in this winter, with a
pulse of 964,600 on November 27, and Amphileptus reaching 14,469 .
on this date and 14,835 a week later. Both species decline during
the flood which follows, and rise during March to culminations, on
the 24th, of 104,535 and 3,636, respectively.
In 1898, Amphileptus disappears on April 12 at 52, save for an
isolated occurrence May 17 at 64. It does not reappear until
October 18 at 52. In 1897, it reappeared October 26 at 59, and
in 1895-96 its limits were 45 and 48, with the exception of one
occurrence, April 17, at 66. Carchesium occurs irregularly and
sparingly during summer months, and Amphileptus was not taken
in the plankton during that period. Its occurrence in the plank-
ton is limited in the main to temperatures below 50, but this
limitation may be due primarily to the reduced numbers, at higher
temperatures, of the organism upon which it feeds. It appears
during the period of greatest sewage-contamination and bacterial
development in the river at Havana. Roux ('01) finds Amphilep-
tus most abundant in stagnant waters about Geneva in the winter
months.
Aspidisca costata (Duj.) Stein. Found in the plankton but once
Jan. 11, 1898, at 32.
Bursaria truncatella O. F. Mull. Average number, 23. This
large ciliate was found in the plankton at irregular intervals and in
119
small numbers. It was found six times in March; twice in January
and April; and once in February, July, and November. Its ap-
pearance in the plankton is thus predominantly in winter months
and at temperatures below 45, though it occurs in the extremes
of temperature conditions.
Carchesium lachmanni S. Kent. Average number, 26,546. This
is normally an attached species, and its appearance in the plankton
is due to the detachment of the heads. Small fragments of colonies
are also found, but the greater number are isolated heads. The
detachment seems to be a physiological process of the organism and
not merely the result of accidents. It is thus a detached and an
adventitious planktont. Many of the heads taken in the plankton
are in a moribund condition. For example, in a pulse of March,
1896, the following proportions were recorded.
Date
Total
Carchesium
per m.s
Per cent,
normal
Per cent,
moribund
1896
March 17
60,420
55
45
" 24
104,535
48
52
" 30
47 571
53
47
April 10
16 688
39
61
Enumerations were based on the total number of heads, both
normal and moribund. The colonies are sessile, and adhere in vast
numbers to any substratum furnishing a suitable place for attach-
ment submerged vegetation, brush, sticks, and fishermen's nets.
The latter sometimes become so clogged with Carchesium and.
floating mats of Crenothrix and Beggiatoa as to break down in the
current of the river. How far the number of free heads in the
plankton is an index of the development of the species in the stream
can not be determined from the data at hand.
This species has been taken in the plankton in every month of
the year, but its occurrences between the early part of May and
120
October 1 that is, above 60 are irregular and the numbers few
(Table I.). It is thus predominantly a cold-water planktont.
Winter collections in 1894-95 and 1896-97 were too few to trace
its seasonal movements. In 1896-97 it appeared November 5, rose
to a maximum of 964,600 on November 27, and declined in the
December- January flood (Pt. I., PI. IX.) almost to extinction, but
recovered during its decline to a minor pulse of 16,160 on January
30. It again fell off in numbers during the floods of February
(Pt. I., PI. X.), but rose during the decline of March to a maximum
of 104,535 on March 17. Numbers become smaller and occurrences
irregular after May 1.
In 1897, Carchesium increased rapidly in late October to a small
pulse of 13,200 on November 2, with a decline in the following fort-
night, and a pulse culminating December 7 at 283,800, with subse-
quent decline. The fluctuations during 1898 may be followed in
Table I. The numbers increase during the slowly rising flood of
January to a maximum of 197,600 on February 8 at 32, and decline
again during the more rapid rise (Pt. I., PI. XII.) of the next three
weeks. Stable conditions in early March bring about a pulse of
89,600 on March 15, and numbers decline again to 2,400 as the flood
passes its maximum in the early part of April. As the levels fall
another pulse of 99,200 appears April 26, from which a descent to
minimum numbers which prevail during the summer takes place
within a fortnight. The floods, especially sudden ones, seem thus
to interfere with the appearance of Carchesium in the plankton,
while gradual rises, as that of November, 1898, are not so detri-
mental.
The table of bacterial occurrences (Jordan, '00) in the Illinois at
Havana and Pekin given on p. 231, Part I., indicates that the bac-
terial development consequent upon the sewage and industrial
wastes of Peoria extends down the river to Havana during the
colder months of the year. The occurrence of Carchesium in the
plankton is thus coincident with that of greatest sewage pollution
and bacterial development at Havana. Carchesium is much more
abundant in the channel of the river, where sewage pollution is
greatest, than it is in the adjacent backwaters. It seems probable
that the bacteria either directly or indirectly contribute towards its
development, constituting, it may be, an important element in its food.
Flood waters, which dilute the sewage (cf . hydrograph and chlorine
121
in PL XLV. of Part I.) might for this reason tend to interfere with
the development of Carchesium, and thus cut off the source from
which the plankton individuals arise. I am not able, however,
to trace any close correlation between the fluctuations of the chem-
ical matters indicative of sewage and sewage decay and those of
Carchesium. In the stable hydrographic conditions of 1897 we find
a symmetrical pulse of considerable dimensions rising from 2,200
on November 9 to 283,800 on December 7, and declining to 26,500
on January 11, 1898. Stable low water with an ice blockade
(Pt. I., PI. XI. and XII.) characterize this season. ~Nr> explanation
for the fluctuation is suggested in the physical environment. The
chemical condition of the water, was, however, greatly disturbed
(Pt. I., PL XLIV.). The fivefold increase in free ammonia is indic-
ative of approaching stagnation under the ice, and the threefold
increase in chlorine marks the sewage concentration. Approaching
stagnation might have caused the decline of Carchesium, or it may be
a specific reproductive cycle of the organism which combines with
the external factors of the environment to produce such a wave of
occurrence.
Chilodon cucullulus Ehrbg. Average number, 102. This species
was found in the plankton in January and February during the bac-
terial increase. It was also found in July. It escapes through the
silk net, and does not ordinarily appear in plankton collections,
though abundant wherever decay is active.
Codonella cratera (Leidy). Average number, 101,024 or 452,500*.
This is the most abundant of the ciliates in our plankton, consti-
tuting about one third of their total number. It appears in
every month of the year, and in 1898 it was recorded in every
collection but one, that of December 13 (Table I.). It is sub-
ject to great fluctuations in numbers, its maximum occurrences tend-
ing to appear in April, May, or June, and again in September or
October. Minimum numbers prevail during the winter, when many
of the shells are empty, and the midsummer interval is subject to
pulses of varying amplitude. Spring pulses were detected as follows :
in 1895, on April 29 (16,324) at 64; in 1896, on April 24 (562,152) at
72;inl897,onApril27(470,000)at60;andinl898,onMay3(736,000)
at 60. These vernal pulses coincide with or approximate closely to
the dates of the spring volumetric pulses. This somewhat remark-
able approximation of dates near the end of April may be the result,
122
in part at least, of the dates of collection ; but after allowance is made
for this, the species still exhibits a seasonal cycle of remarkable regu-
larity. The autumnal pulse is of less amplitude, and of less regu-
larity in location as to time and temperature. In 1894 it appears
September 4 (14,000) at 78; in 1895, on September 12 (5,840) at
81; in 1896, on August 29 (58,800) at 74 or October 14 (63,200)
at 57; in 1897, on October 5 (204,400) at 71; and in 1898, on
September 27 (92,800) at 73.
The midsummer pulses are, as a rule (Table I.), of less amplitude
than the vernal or autumnal ones. In 1896 and 1898 exceptions to
this statement appear in two large developments which follow in
each case upon the decline of the June rise. In 1896 (Pt. I., PI. X.)
this pulse (152,400) came June 11, and in 1898 (Pt. I., PI. XII.) it
came (1,499,200) June 7 at 78 and exceeded in amplitude the re-
corded vernal pulse. In both cases the pulse was recorded as occur-
ring at an interval of a week after the crest of the June rise had
passed. The character and sequence of these pulses is well shown in
Table I.
The occurrence of Codonella in abundance in the purer backwaters
and in the plankton of our Great Lakes (Kofoid, '95) indicates that it
is not dependent upon the sewage bacteria directly for food for its
development in our waters. The appearance of the greatest pulses
during a period of considerable sewage dilution still further indicates
its independence of sewage bacteria. A comparison of the fluctua-
tions of the totals of the chlorophyll-bearing organisms with those of
Codonella affords some evidence of a correlation between the two.
Of 39 pulses which can be traced, in our records in the chlorophyll-
bearing organisms, 21 precede and 13 coincide with those of Codo-
nella, while in the remaining 5 instances the multiplication of Codo-
nella precedes that of the phytoplankton as a whole. Thus in the
main the pulses of Codonella follow, or coincide with, those of the
phytoplankton. The evidence of this sequence may be followed in
Table I. by a comparison of the records of Codonella with those of the
total phytoplankton. The sequence indicates that the food of Codo-
nella may be found in the phytoplankton, and that these recurrent
periods of growth have some connection with the conditions of nu-
trition. The seasonal cycle of Codonella is closely followed by the
other member of the family found in our plankton Tintinnidium
fluviatile.
123
Codonella occurs throughout the whole range of temperatures. The
winter minimum and the decline during the maximum temperatures
of summer, combined with the presence of vernal and autumnal, or
late summer, pulses, indicate that the optimum conditions for this
organism lie neither in winter nor in summer. The spring pulse
was at temperatures of 60-72, and the autumnal one at a wider
range of 57-78. Permanent increase in numbers does not begin
(Table I.) until March 15 at 46, and the permanent falling off is
found on November 15 at 41. The optimum temperatures in our
waters thus lie near 60-70, and conditions favoring growth are
limited to a range of 10-15 upon either side of the optimum.
This species readily escapes through the silk net on account of its
small size and its motility, and such collections give at the best in-
complete evidence of its seasonal distribution. The amplitude of its
fluctuations is thus reduced, and owing to the irregularity of the
error arising from leakage, the reduction is not proportionally distrib-
uted throughout the year. Tests made of the loss of Codonella by
leakage through the silk indicated that but one was retained to
twenty-four found in the filtrate . Codonella was counted in both the silk
and filter-paper collections, with the result that in 1897 the totals for
the year (omitting one date on which the filter collection contained
an unusually large number of Codonella) showed one Codonella in the
silk to twenty-five in the filter collection. In 1898, however, the
ratio was one to four and a half. The error in the filter collection
is large, but data seem to justify the conclusion that only a small
proportion of the Codonella is retained within the silk net. The
proportion for the whole period of collection by the two methods
(August 3, '97, to March 28, '99) is one to seven, if one date on which
aberrantly large numbers appear in the filter collections be omitted.
This species is a typical planktont, and is apparently the same as
C. lacustris Entz, by which name it is designated by European writers.
Leidy's name, however, has priority according to the accepted rules
of nomenclature. It is an exceedingly variable organism, at least
in the form, proportions, and size of the shell, in the degree of its con-
striction, and in the foreign particles which fill its matrix. The rings
or bands which ornament the orifice vary in their number, width, and
relative proportions, and in the perfection of their development.
The intergradation which these variants exhibit is sufficient to my
mind to make their elevation to specific rank unjustifiable.
124
Codonella is an important element in the food of many of the lim-
netic rotifers, especially Asplanchna.
Codonella is a common constituent in the plankton of our own
Great Lakes (Smith, '94; Kofoid, '95; Jennings, 'OOa), and has
been reported from most European waters. Apstein ('96) finds in
German lakes major pulses in spring and autumn and minor ones in
midsummer. Lauterborn ('94) reports Codonella in the plankton of
the Rhine, and Schorler ('00) in that of the Elbe, but neither follows
its seasonal history.
Coleps hirtus Ehrbg. Average number, 1 3 . This species occurred
in the plankton collections irregularly and in small numbers, princi-
pally in autumn months during the height of the bacterial develop-
ment. It escapes through the silk readily.
Colpoda cucullus Ehrbg*. Average number, 9,615. This species
appears in the plankton principally during the colder months of
bacterial predominance, from November to April, and occasionally
during the summer.
Cotkurniopsis vaga (Schrk.) Blochmann was found in both 1898
and 1899 on Canthocamptus.
Didinium nasutum (O. F. Miill.) Stein*. Average number,
12,692. This species also is found in the plankton during winter
months, especially in November and December during the bacterial
increase. It was also found in midsummer.
Epistylis spp. Average number, 2,020. The free heads or frag-
ments of colonies of one, or possibly of several, unidentified species of
Epistylis, or it may be of Opercularia also, were associated with Car-
chesium lachmanni in the plankton during the colder months, but in
much smaller numbers (1 to 13 in 1898). Identification in most cases
was impracticable, though in some instances E. flavicans Ehrbg. was
determined, and it seems probable that most of the winter forms at
least belong to this species. Hempel ('99) reports E. plicatilis on
snails, and various other aquatic animals have been found infested
with colonies of undetermined species of Epistylis.
The distribution of Epistylis in the plankton (Table I.) is in its
limits somewhat like that of Carchesium. It is more abundant and
more continuously present during the period from November to June
(at temperatures below 60) than in the intervening warmer months.
It is found throughout the whole range of temperatures. Its pulses
coincide with those of Carchesium when they occur, but they are not
125
always found in Epistylis when they appear in Carchesium. This
degree of similarity in the seasonal cycle of the two genera is indica-
tive of their correlation with the same environmental factors, the
principal one of which is the increase in bacteria attending the colder
months.
Euplotes char on (O. F. Mull.) Ehrbg. was taken but once in the
plankton August 23, 1898.
Euplotes patella Ehrbg*. Average number, 2,888. It was found
in small numbers and at irregular intervals from April to December
throughout the full range of temperatures. It was most frequently
taken in the summer.
Glaucoma scintillans Ehrbg.* Average number, 39,615. This
species was taken in the plankton from the middle of October till the
middle of April. It was present in larger numbers and more contin-
uously in December and February. It is thus a member of the
plankton during the time of bacterial increase.
Halteria grandinella O. F. Mull.* Average number, 255,769.
The seasonal distribution of this species in the plankton does not
show the limitation to the winter months noted so frequently in other
ciliates. It was found in every month of the year but May, in largest
numbers in July and August, and most continuously in December and
January. The data are too few and irregular to determine any pre-
dominance as to season or temperature.
Holophrya simplex Schew. was found in small numbers in the
filter collections of December, February, and March in the winter of
1896-97 at temperatures from 32 to 44.
Leucophrydium putrinum Roux. Average number, 525. This
species was recorded July -September, 1898, during the low-water
period, at temperatures from 89 to 63. It was described by Roux
('99) from stagnant water, but in our plankton no conditions of stag-
nation attend its presence, though sewage contamination is great and
decaying organic matter abundant.
Lionotus spp. Average number, 94. With Amphileptus in the
winter plankton there occur a number of other, smaller, gymnostome
ciliates which in best-preserved specimens resemble Lionotus. A few
occurring in March and April, 1898, were found to be L. fasciola
Ehrbg. , and it is probable that most of the individuals belong to this
species, though exact identification is difficult with plankton mate-
rial. The seasonal distribution of Lionotus coincides very closely
126
with that of Amphileptus. The species appear in November or De-
cember and continue through March in temperatures below. 50, but
the numbers retained by the silk net are too small to trace their sea-
sonal routine. Their seasonal distribution in the plankton "coincides
with the period of greatest access of sewage and bacterial increase in
the river at Havana. Roux ('01) finds this genus well represented in
the fauna of swamps, and most abundant in October and March.
Loxodes rostrum Ehrbg. was identified but once March 22, 1897,
at 44.
Nassula ntbens Perty occurred July 30, 1897, at 84.
Opercularia articulata Goldf. This species is parasitic upon
aquatic Coleoptera. In the plankton of June 28, 1897, eleven
colonies or fragments of a colony were found, the largest with 115
zooids.
Opercularia nutans (Ehrbg.). Average number of zooids, 60.
In the plankton this species was found attached to Alona afjinis in
January, 1898, and to Cyclops in April and August.
Opercularia not specifically determined were found free in the
plankton in June and July ; in November, attached to Canihocamptus ;
in January, attached to Brachionus and even to the eggs of this
species. An unidentified form was also found upon Cyclops.
Ophryoglena atra Lieberk. Five irregular occurrences of this
species in small numbers were recorded in 1899 from January to the
middle of March.
Paramecium spp. Average number, 41. Paramecium was
found 18 times in the plankton. Two of these instances were in May
and August at temperatures of 64 and 79, and the remainder were
between November 20 and March 30 at temperatures below 48.
Most of the occurrences are in midwinter at minimum temperatures
under the ice. P. aurelia (O. F. Mull.) has been found in the river
waters (Hempel, '99), but not all taken in the plankton belong to this
species. Specific determinations are not easily made with accuracy
in preserved plankton material. In our plankton, Paramecium is
present principally during the period of greatest contamination by
sewage.
Plagiopyla nasuta Stein*. Average number, 1,181,000 during
the winter of 1898-99 from November 29 to March 28. This species
was not recognized in the plankton of previous winters. It reaches
a pulse of 11,520,000 on January 3, 1899, at 32.2 under the ice.
127
Levander ('94) finds it in numbers under the ice in Finnish waters.
On account of its motility and small size it readily escapes through
the silk net.
Pleuronema chrysalis (Ehrbg.) Stein. Average number, 9. Re-
corded only in January, 1898, at minimum temperatures.
Prorodon farctus Clap, and Lach. Only a few scattered occur-
rences from the last of September to the first of March at tempera-
tures from 73 to minimum. An unidentified species of Prorodon was
also found irregularly from November to April.
Pyxicola affinis S. Kent. Average number, 58. This species is
usually attached to aquatic plants, especially to Lemna. It has been
found in the summer plankton from June to August during maximum
temperatures, especially in 1896, when recurrent floods brought much
Lemna from the backwaters into the river. It was found October 18
at 52, attached to Melosira varians.
Rhabdostyla spp. Average number, 1 10. Peritrichan ciliates re-
ferred to this genus have been noted on Cyclops, Canthocamptus,
Oligoch&ta, and even in considerable numbers upon the body, append-
ages, and eggs of Polyarthra platyptera. They have appeared thus
passively in the plankton during winter months from December to
March, especially in 1899.
Stentor casruleus Ehrbg. Average number, 882. This species
presents a characteristic seasonal distribution in our plankton. Its
numbers are never very large, and its full cycle can not always be
traced in the records. It is a planktont of the colder season in our
waters. But three records one July 28, 1896, at 82, one August 3
of the same year at 80, and a third, August 15, 1894, at 84 lie
outside of the period between September 1 and May 1. In 1898
(Table I.) the autumn cycle begins September 6 at 79, but in
both 1895 and 1897 the species does not appear until late in
November or in December at 34 or below. In years prior to 1898
the numbers were small and irregular, but on January 21, 1898,
the maximum number of 28,800 was reached at 34, under the ice,
during the slowly rising flood of that month (Pt. I., PI. XII.). It
accompanied an increase in Stentor niger, and there are indications
elsewhere that the two species may fluctuate together. The high
(Pt. I., PI. XLV.) chlorine (38.), nitrites (.175), and free ammonia
(4.6) at the season of greatest development in the plankton are in-
dicative of conditions approaching stagnation. The appearance of
128
this species in stagnant water has often been observed. Roux ('01)
finds it especially abundant in September, October, and February in
stagnant waters about Geneva.
Stentor niger Ehrbg. Average number, 3,124. In our waters
this species also is a winter planktont (Table I.). There have been
but four records of occurrence between May 1 and September 1 . In
1895-96 the species appeared November 14 at 44 and reached a
maximum of 68,635 December 18, after three weeks of minimum
temperatures and approaching stagnation under the ice. Numbers
declined in the December- January flood (Pt. I., PI. X.), but rose
again in March, as the flood declined, to 39,087 on the 24th at 40.
It disappeared from the plankton April 30 at 70 and did not re-
appear until November 1 7 , from which time it continued until March
22. In 1897-98 it returned September 21 at 71, attained a maxi-
mum of 42,000 November 23 at 43, declined during December, and
rose to 47,000 on January 21 at 34 under the ice, and in the con-
ditions approaching stagnation described in connection with the dis-
cussion of S. c&ruleus. A decline in numbers continued until April
12 at 52. Favorable conditions for growth are thus found in our
waters between 32 and 50, and the optimum seems to lie near 40
or below.
This species reaches its greatest development in our waters during
the time of greatest sewage pollution and bacterial development. It
is known as a bog-water species, and was found by Roux ('01) in
stagnant waters about Geneva during the colder months. Hempel
('99) reports this species as 5. igneus (?), but from the descriptions
of Roux ('01) I am inclined to consider it as 5. niger Ehrbg. It may
be that both species are included in our data, but they are predomi-
nantly of the niger type. They include also individuals of the black-
ish variety 5. igneus var. juliginosus Forbes, which, it would seem
from Roux's description of these species, should be transferred to 5.
niger. The fuliginosus form was very abundant in the margins of
Pine and Round lakes, Michigan (Kofoid, '95), during the summer
in surface temperatures of 61-70, where sewage contamination was
but slight.
Stentor polymorphus (O. F. Mull.) Ehrbg. was found sparingly in
July and August during maximum temperatures. Hempel ('99)
reports S. barretti Barrett and 5. roeselii Ehrbg. from the river, but
I have not identified them in the plankton collections.
129
Strombidium viride Stein was found in small numbers in January-
March, 1899, at minimum temperatures.
Stylonychia mytilus (O. F. Mull.) Ehrbg. was found in the plankton
sparingly from September to February, and once in June.
Tintinnidium fluviatile Stein. Average number, 22,590 or 1,640,-
192*. This species is somewhat sharply limited to the warmer
months in its seasonal distribution. In 1898 (Table I.) it makes its
appearance April 4 at 49, reaches a maximum of 720,000 May 3 at
60, and has three decreasing pulses ; one of 1 04, 000^ oa June 14 at 80,
one of 95,200 on August 2 at 79, and one of 22,400 on September 27
at 73, and disappears from the plankton October 18 at 52. The
records in previous years are more irregular, though traces of vernal
and midsummer pulses can be found in the records. Filter-paper
catches indicate that only one in eighty of this species is retained by
the silk. They also locate the pulses as approximately coincident
with those of the silk collections.
Apstein ('96) finds Tintinnidium to be a spring planktont with
its maximum in April in Lake Plon, while Seligo ('00) finds it in lakes
near Danzig in the autumn, with a maximum in September. In our
own waters in 1896 the autumnal pulse in August-September exceeds
the vernal one.
The gelatinous lorica of this species is subject to great variation in
its size and proportions, and especially in the region about the aper-
ture. A somewhat thimble-shaped form was described by Hempel
('96) as T.illinoisensis, the specific distinctions being based wholly on
the lorica. This form intergrades with the typical lorica of T.
fluviatile Stein, and should not in my opinion be given specific rank.
Trachelius ovum Ehrbg. Average number in 1895, 847. This
species did not occur in 1898 but was rather common in November-
December, 1895, reaching a maximum of 10,695 on December 4 at
32.5. Isolated appearances in small numbers in December and
January of other years have been recorded. In our waters it is thus
a winter planktont. Stagnation conditions under the ice were
approaching (Pt. I., PI. XLIII.) when the pulse of 1895 occurred in
the Illinois River. Apstein ('96) found it, however, in Lake Plon with
a maximum in May- June, disappearing in the summer and returning
again in November.
Trichodina pediculus Ehrbg. Average number, 1; in 1897, 874.
This species is normally found upon Hydra, on the gills and skin of
130
amphibians, and on young fish. It appears in the plankton during the
summer months in every year except 1898, a single record only being
made in that year. The earliest record was on June 1 1 , and the latest
on November 3 1 . The whole temperature range is practically included
in these occurrences, though the species disappears within a few weeks
after the temperature falls below 50. It usually appears in small
numbers and irregularly, and no pulses like those of typical plank-
tonts can be traced. A free life in the plankton is apparently not its
usual habit. Zacharias ('00) has recently called attention to its
appearance in the plankton in German waters.
Vorticella rhabdostyloides Kell. Average number, 61. This little
Vorticella is found attached in small clusters to Anabozna spir aides
and occasionally to other members of the phytoplankton. It is some-
what common in the waters of Lake Michigan, but is rare in spring
months in the Illinois River.
Vorticella spp. Average number, 7,843. At irregular intervals
from April to November isolated individuals and small clusters at-
tached to bits of debris in the silt were taken in the plankton. They
were most abundant at temperatures above 50. The irregularity
in their occurrences indicates that they are adventitious in the plank-
ton. Identifications of plankton material are impracticable except in
strongly marked species. Hempel ('99) has found V. campanula
Ehrbg., V. microstoma Ehrbg., and V. similis Stokes in the river and
its adjacent waters.
Zoothamnium arbuscula Ehrbg. A few colonies were taken in
August and September in 1896 in the plankton, probably adventitious
during the disturbed hydrograph of that year (Pt. I., PL X.).
The preceding list of 45 species does not complete the catalog of
the ciliate constituents of the plankton, though it includes all of the
species of quantitative importance during the years of our operations.
The residium of unidentified ciliates, which, excluding the partial
identifications in the above list, does not often exceed two per cent,
of the total individual ciliates, includes principally isolated individ-
uals of species difficult of identification or others whose preservation
did not permit it, and a considerable number of small ciliates and of
forms ectoparasitic upon Entomostraca and other planktonts. Most
of these organisms are either adventitious or passive members of the
plankton, and further study of the littoral region, of stagnating
131
waters, and of these parasitic forms will reveal the great richness of
the ciliate fauna in this aquatic environment.
SUCTORIA.
Average number, 332. This class is not quantitatively im-
portant in the plankton, being represented, in so far as our records
go, only by adventitious or passive planktonts. No limnetic species
has as yet been found in the Illinois. An examination of the
littoral region during the prevalence of ciliates wilt probably yield
a rich suctorian fauna.
DISCUSSION OF SPECIES OF SUCTORIA.
Acineta linguifera Clap, and Lach. This species is usually found
on aquatic Coleoptera. A single occurrence of an unattached indi-
vidual was recorded June 21, 1898.
Metacineta mystacina Ehrbg. Average number, 301. This
species occurred in the plankton from March till October in 1898 and
in the winter months of 1899, at irregular intervals and in small
numbers (Table I.). Most of its occurrences attend flood invasions,
and it is evidently adventitious. It is frequently attached in the
plankton to minute particles of debris. This species varies greatly
in the size of the lorica. Sand ('01) gives the range in height as
from 3 3-7 00 -/i. The variation in proportions has given rise to a
number of descriptions of new species by Stokes ('88 and '94) and
Maskell ('87), but an examination of a series of individuals such as
appear in the plankton shows that they intergrade so closely that
specific distinctions can not be maintained for the variants. Meta-
cineta appears throughout the whole range of temperatures, no
seasonal predominance appearing in the records.
Podophrya fixa O. F. Mull. Average number, 12. This species
is also adventitious in the plankton. It was recorded in March and
September at 37 and 73. Cysts were noted January 21.
Tokophrya quadripartite Clap, and Lach. Average number, 4.
Adventitious in the plankton in March and November. Hempel
('99) finds it most abundant in May and June, associated with
Epistylis plicatilis and Opercularia irritabilis on crayfish, insect
larvae, and turtles.
Tokophrya cydopum Clap, and Lach. Found occasionally upon
Cyclops during spring and summer.
(10)
132
PORIFERA.
Spongilla spp. Average number of spicules, 772. The identifi-
cation of fresh-water sponges by isolated spicules is practically
impossible, and, moreover, the sponge fauna of the Illinois River is
as yet practically unknown. No attempt, therefore, was made to
identify the species to which the spicules which occur in our plank-
ton collections belong. They belong to the genus Spongilla in part,
and were usually the simple sarcode forms, the gemmules or their
spicules not appearing in the plankton. They occurred in all
months of the year, and were found in 46 per cent, of the collections.
They are adventitious, and their occurrence in the plankton is there-
fore dependent in part upon hydrographic conditions. Records in
December and January are few (3) and always occur on rising floods.
In February and March, months of rising floods, they are increased
(8 and 7), but decline again in April- June (3,5, and 5), months of
predominantly declining water and more stable conditions. In
midsummer and autumn months (July to November) they again
occur more frequently (8 to 12), probably as a result of proximity
to the season of greatest growth and frequency of sponges in the
river and its backwaters. Here also they occur most frequently
in years of greatest hydrographic disturbance, as, for example, in
1898. The adventitious relation which they bear to the plankton
is also seen in their erratic and irregular numbers. The maximum
record (16,000 per m. 3 ) was made June 28, 1897, on the rising flood;
the next in size, on August 10 in stable low water. In both instances
the plankton was probably taken from water in which as a result of
some local disturbance the remains of some disintegrating sponge
had been distributed. Living sponges are found in considerable
abundance on submerged brush and timbers in the channel and
backwaters during the summer months, and feed on the smaller
organisms of the plankton, being one of its depleting agencies.
CCELENTERATA.
Hydra fusca L. Average number, 39. Hydra occurred in about
16 per cent, of our channel collections a percentage which would
be considerably increased if the whole of each collection had been
examined for it, or if backwater collections should be included. With
one exception the 28 occurrences recorded, all fall in May-September
133
at temperatures rarely below 70. The earliest record in channel
waters was on May 1, 1896, at 68.75, and the latest on November
15, 1897, at 47. Of the 28 records in channel waters the months
from May to September have, respectively, 6, 3, 10, 7, and 1 record,
and there is 1 in November. Hydra is thus a late vernal and a
summer planktont in our waters.
Observations in the field and a cursory examination of the col-
lections made in the backwaters have indicated that Hydra is often
very abundant on the vegetation. It is also limnejtic in habit,
floating with the foot attached to the surface film and tentacles
widely extended; or, without attachment, in the deeper strata of
water. A similar limnetic habit was often observed in the case of
Hydra in channel waters, especially on still warm days when the
surface was unruffled.
Hydra \vas generally more abundant in the plankton in May or
in early summer. The maximum record in channel waters was
3,200 per m. 3 on July 21, 1897, the error of dilution being, however,
large in this record. In Quiver Lake on May 8, 1896, a maximum
record of 5,335 per m. 3 was made, the error of dilution being very
small. This was during a vernal plankton pulse (8.14 cm. 3 per m. 3 )
in these waters, when the food of Hydra was present in considerable
abundance.
Hydra viridis L. was seen 'frequently in spring-fed backwaters
and in laboratory aquaria, but was never recognized in plankton
collections made in channel or backwaters. The limnetic habit
noted in H. fusca was not observed in the case of this species. .
PLATYHELMINTHES.
TURBELLARIA.
Numerically and from the volumetric standpoint the Turbellana
are not of great significance in the plankton of fresh waters as a rule.
However, in some seasons and under certain conditions Stenostoma
becomes very abundant, as, for example, in autumn months in back-
waters, and generally where decaying vegetation abounds. In the
autumn of 1895 the plankton in the relict pools of Flag Lake consisted
almost entirely of Synura uvella, Stenostoma leucops, and Entomos-
traca.
134
The average number in channel waters is 103 per m. 3 , and, as
might be expected, their occurrences are erratic in seasonal distri-
bution and their numbers are irregular. They occurred in channel
waters in eveiy month of the year and throughout the whole seasonal
range in temperatures. The numbers in 1898 were larger and occur-
rences more frequent in May, during the run-off of the spring flood,
and smaller and more erratic during the rest of the year. In the
total of all collections enumerated the percentage of occurrences was
highest in June (60 per cent.), July (83 per cent.), August (48 per
cent.), and October (47 per cent.), and lowest in colder months, when
it rarely rises above 30 per cent. The numbers are also larger in the
warmer months, a maximum record of 19,250 per m. 3 on September
4, 1894, following a slight rise in river levels at low stages. The
adventitious character of the Turbellaria in channel plankton is sug-
gested by the erratic data, but the adaptability, at least of certain
species, to the limnetic habit under certain conditions is also indi-
cated by the large numbers.
The identification of the Turbellaria in plankton collections is not
feasible in the course of the usual methods of examination of pre-
served plankton. Accordingly no effort was made to identify the
individuals occurring in our catches. Many of them were evidently
rhabdoccele turbellarians, and of these probably many were Stenos-
toma leucops. The genus Vortex was also represented.
Mesostomum ehrenbergii O. Schmidt was taken in small numbers
on August 26, 1895, along the shores of the river in vegetation. This
identification is that of Dr. W. McM. Woodworth ('97).
Stenostoma leucops O. Schmidt. Average number, 21. By far the
greater proportion of the turbellarians in our collections probably
belong to this species. The statements made regarding the group as
a whole therefore probably apply to this species.
TREMATODA.
Many of our predaceous fishes and other aquatic vertebrates are
infested to an extraordinary degree by flukes parasitic in the intestine
or other viscera. This, in conjunction with the fact that the fish
markets are located in house-boats along the stream and their refuse
generally cast directly into the channel, is sufficient to account for the
few adventitious adult distomes which have been noted in our plank-
135
ton collections. They have occurred singly in February and July,
but were not identified.
The free-swimming larval stages or cercaria of unidentified trem-
atodes were also found singly in August, September, and October.
Aspidogaster conchicola v. Baer, which occurs abundantly in the
mantle cavity and pericardium of many of the Unionidcz (see Kelly,
'99), which form great beds on the river bottom, was taken in an
immature condition in the plankton on June 27.
Cotylaspis insignis Leidy, likewise a parasite of the Unionidce,
associated with Aspidogaster but confined principally to the mantle
chamber, was taken in the plankton on February 4.
CESTODA.
Tetrarhynchus sp. was -adventitious in the plankton on June 27,
and doubtless of similar origin to the adult trematodes above noted.
NEMERTINI.
Fresh-water nemerteans were definitely identified as such in the
plankton on only two occasions, July 23, 1894, and March 22, 1897.
They were doubtless adventitious from the shore or bottom, where
they are most abundant.
NEMATELMINTHES.
NEMATODA.
The free-living nematode worms are predominantly shore and
bottom forms, living in the midst of the decaying organic matter of
the bottom ooze. In a habitat such as ours, where the quantity of
this decaying matter is very great, the nematodes are correspondingly
abundant, and, owing to the unstable hydrographic conditions, they
find many opportunities of joining the plankton temporarily. Ac-
cordingly we find that nematodes are met most frequently and in
largest numbers in rising flood waters, when the bottom deposits of
tributaries and the main stream are carried in channel waters as silt.
Thus, in the month of March nematodes occurred in 13 of the 15
collections examined, with an average number per m. 3 of 465, while
in August they were found in but 8 of 21 collections, and averaged
only 186 per m. 3 . So, also, in the winter flood of 1895-96 nematodes
were found in the plankton almost continuously till the middle of
136
April, while in the more stable conditions of the preceding year they
were found in only one third of the collections. In 1897 most of the
31 collections examined were made in stable conditions, and nema-
todes were found in but 5 of these, and 4 of these 5 were made in
rising flood waters. In 1898, a- year of greater hydrographic dis-
turbance, nematodes occurred in 31 of the 52 collections, averaging
318 per m. 3 to 82 in 1897. Of the 31 occurrences in 1898 all but 6
were in recent flood waters. The hydrographic conditions attending
the presence of nematodes in the plankton thus indicate that they are
adventitious in the plankton. Further evidence of this is to be found
in their erratic numbers. Thus, on February 20, 1896, none was re-
corded, and on the 25th their numbers rose in flood waters to the
maximum record for all of our collections 18,422 per m. 3
No effort was made to determine the species of these nematodes.
A considerable variety of forms awaits the labors of some courageous
systematist.
ACANTHOCEPHALA.
These worms are found abundantly in the Catostomida and other
limophagous fishes of the Illinois River, and in many of the water-
fowl which feed in its waters. A chance occurrence of a single
specimen in the plankton on August 3, 1896, is probably to be ac-
counted for as in the case of other intestinal parasites.
ANNULATA.
OLIGOCH.
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146
147
the seasonal distribution of individual species and these recurrent
plant pulses will be discussed in connection with the various species
wherever the data are available. For the present it will suffice to call
attention to such correlation as exists between fluctuations of the
phytoplankton and the total Ploima. The table on the preceding page
gives the location and amplitude of the maxima of the ploiman pulses,
and a graphic presentation of the seasonal curve of distribution of the
total Rotifera will be found in Plates III. and IV. On comparison of
the ploiman pulses with those of the chlorophyll-bearing organisms,
graphically presented in Plates I. and II., it will be found that 15 of
the 33 pulses of Ploima contained within the period covered by the
plates coincide in location with the plant pulses ; that 12 follow at the
next collection, usually a week later, and 3 within a fortnight ; while
only 3 of the 33 exhibit no such correlation. The data suggest
strongly the agency of the plant pulses in building up the Ploima,
and that the food relations are fundamental in the fluctuations of
these planktonts.
DISCUSSION OF SPECIES OF PLOIMA.
Anuraza aculeata Ehrbg. Average number, 1,839. In 1898
this species has a very well-defined and characteristic seasonal
distribution (Table I.). It first appears March 8 at 37, increases
to a maximum of 45,200 on May 10 at 61. then declines, and
disappears June 14 at 83. The curve of its occurrence in this year
is a very symmetrical one. It reappears on December 27 at 32, and
there are scattered occurrences through the winter months of 1899.
Records in other years suggest in the main a similar distribution.
In 1896 it first appeared January 6, rose to a pulse of 6,550 on May 8
at 76, and, on the decline of the June rise, there was a second and
larger pulse of 29,600 on June 17 at 76. It reappeared on Decem-
ber 29, and in 1897 reached a vernal maximum of 22,400 on May 25
at 66, then disappeared, and was not again noted in the following
winter nor until March 8. In 1894 the last vernal record was made
June 12, and on September 4, at 78, there was an autumnal pulse
of 13,825 a phenomenon not repeated in subsequent years. The
normal course of its seasonal distribution in the river plankton seems
to be as follows: reappearance in December when minimum
temperatures have been reached; slow multiplication during the
winter, and a well-defined pulse on the decline of the spring flood in
(in
April-May with the possibility of a second on the June rise; and
prompt and complete disappearance when maximum summer tem-
peratures are established. Low water in the autumn seems to inter-
fere with an autumnal pulse. In 1894 there was a well-sustained
rise in September (Pt. I., PL VIII.) and a pulse of A. aculeata. In
1896, however, no pulse occurred in the high water of the autumn.
No midwinter occurrences followed the very low water of 1897. It
is thus in channel waters a vernal planktont, with its temperature
optimum near 70 but below the summer maximum. Hempel's
statement ('99) that it is a "winter species" is borne out by its
presence from December through the winte'r, but its numerical
distribution ranks it at once with the vernal organisms. Lauterborn
('94) finds it abundantly in winter months in the Rhine, and Ap-
stein ('96) speaks of it as a "Sommerform," absent from Lake Plon
from November till March, and with maxima from April to July in
different bodies of water where it continues through the summer
and till October, and then disappears. Summer temperatures in
these waters, however, are not recorded by him above 21 C. (69.8
F.), which is about the temperature at the time of the vernal maxi-
mum in the. Illinois, and at least 10 F. below that of the summer
maximum in our waters. Jennings ('94, '96, and '00) records it as
abundant in the summer plankton of Lake Erie, Lake Michigan, and
some inland lakes of Michigan. These waters also are somewhat
cooler (5-10 F.) than those of the Illinois River in midsummer.
Temperature, it seems, must have a decided effect upon the seasonal
distribution of this organism in our waters, though the chemical
conditions and food supply may also enter as factors in the summer
suppression of the species.
Females carrying usually a single egg appeared in 1898 early in
April, and were most abundant during the maximum of the pulse.
On an average, less than a fourth of the females were ovigerous.
Empty loricae appeared May 10 (4,800) and 17 (3,200) at the crest
and decline of the spring pulse, and the same phenomenon of deca-
dence was noted in previous years during this period. Outbreaks
of parasites were not recorded for the species, and the decline is to
be attributed to cessation of reproduction and to the death and
destruction of the individuals by the more usual causes.
This species is quite 'variable, but no effort was made to follow
its seasonal history. The type form is by far the most abundant.
149
A. aculeata var. valga Ehrbg. was seen frequently. A. serrulata
Ehrbg., regarded by Weber ('98) as a variety of A. aculeata, was
recorded Jan. 24, 1899, and found by Hempel ('99) in December. It
seems to be rare in our plankton. Forms approaching A. aculeata
var. brevispina Gosse were also noted, but they, too, are rare, being
recorded only in February and March, 1899. A. aculeata var.
curvicornis Ehrbg. was noted April 29, 1896, at 70.
Anurcea cochlearis Gosse. Average number, 69,393, distributed
as follows: A. cochlearis (sensu strictu) together with A. cochlearis
var. macracantha Lauterborn, 9,421; A. cochlearis var. tecta Gosse,
15,432; and forms with posterior spine of intermediate length
between cochlearis and tecta which include A. cochlearis var. stipitata
Ehrbg., 44,540. Numerically this is one of our important species,
containing over one ninth of all the rotifers in 1898. It is surpassed
only by Brachionus bakeri (with varieties included) , Polyarthra, and
SynchcBta. Average number of eggs, 32,358.
This is a perennial planktont, appearing in every month of the
year throughout the whole range of temperature. Its entire absence
in August, 1898 (Table I.), is not paralleled in any other year. In
1897, for example, there is a well-developed pulse of 45,600 on
August 24. In 1894, 1895 , and 1896 there is a midsummer minimum
of a few weeks' duration in July, August, or September, but it is
irregular in its location.
While the appearance of sexual cycles was not traced by the
records of males and winter eggs, a matter of some difficulty and
uncertainty in preserved plankton material, the existence of such
cycles is suggested by the recurrent pulses of occurrence in this
species (Table I.). It is possible that the species is poly cyclic in
our waters. The pulses in 1898 are well defined, in fact, somewhat
better than in previous years. The following table gives the num-
bers in the pulses in the several years and the dates and tempera-
tures at which the maxima occurred.
All of the large pulses save those of November and December
and one at the close of October (Oct. 25, 1898, 28,500) lie at tempera-
tures above 60. The vernal pulse of April-May is the largest and
appears between 60 and 70, and the amplitude diminishes as the
period of maximum heat progresses, though in 1898 there was a
recurrence of larger numbers as temperatures fell. The optimum
150
PULSES OF ANUR^EA COCHLEARIS.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1896
May 8
76
100,870
June 11
73
95,200
July 2
28
81
81
12,800
17,600
1898
May 10
62
1,145,600
June 21
77
372,800
July 19
84
17,200
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1894
1895
Sept. 4
Sept. 23
78
76
7,350
1,521
Aug. 21
83
17,805
Nov. 20
44
1,120
1896
1897
1898
Aug. 21
Aug. 24
79
78
5,600
45,600
Sept. 16
Oct. 5
Sept. 27
Oct. 25
71
70
73
48
6,224
4,800
54,400
28,500
Dec. 29
35
3,840
Nov. 21
40
10,000
conditions seem thus to be found in the river at temperatures some-
what below the maximum, between 60 and 70.
The phenomena of recurrent pulses are distinctly traceable in the
seasonal distribution of this species, not only in 1898 (Table I) but
also in preceding years. The large May and June pulses of 1898
appear on the declines of the spring and the June rise, respectively;
the pulse of September 27 is in a falling river; and that of October
25, on a slowly rising flood (Pt. I., PI. XII.). In 1897 (Pt. L, PL
XI.) the first two pulses attend the spring flood and June rise in like
manner, but the two subsequent pulses are in stable low water. In
1896 five of the seven pulses lie on the declines of the recurrent
floods of that year and two in rising waters (cf. PI. X. of Pt. I. and
the table just given). In 1894 and 1895 the pulses appear either
in falling water or in the earliest stages of the rise. The number of
pulses on declining waters is somewhat greater than the relative
number of days of this condition w 7 ould lead us to expect, and it
seems probable that optimum conditions for the appearance of
larger numbers of Anuraza cochlearis are to be found in such hydro-
graphic conditions. The run-off of impounded backwaters is one
of the favorable phases during flood decline. On the other hand.
151
the distribution of the pulses with reference to the floods and the
appearance of pulses during rising water suggest the operation of
other factors than the one arising from contribution from back-
waters.
The pulse must be dependent to a large extent upon food supply
of the organism, and a correlation between its periods of multiplica-
tion and the pulses of its food, the chlorophyll-bearing organisms, is
to be expected. A comparison of the seasonal distribution in 1898
(Table I.) and the pulses of chlorophyll-bearing organisms (PL II.)
reveals the fact that three of the A. cockle aris pulses coincide with
those of the plants constituting their food, and the other three
coincide in part only, the remainder of the chlorophyll-bearing
groups reaching their culmination a week prior to that of the rotifer.
In 1897 the three pulses of A. cochlearis which lie in the common
period (PI. II.) all culminate a week (in one case in part in fourteen
days) after the maximum of the plants in question. In 1896, three
pulses coincide and three follow in the subsequent collection; and
in 1895, two coincide and two follow. Collections at daily intervals
would be necessary to follow the correlation more accurately. It is
probable from these juxtapositions and sequences in the A.
cochlearis-algse pulses that we are dealing with a food relation.
Multiplication of algae leads to increase of Anur&a, which, in turn,
reduces the algae, and then itself declines until the food planktonts
again increase.
Anurcsa cochlearis is exceedingly variable in the length of the
posterior spine, in the development and degree of curvature of the
anterior spines, in the arrangement of the areas of the lorica, and in
the degree of its ornamentation by small spinules. The separation
of these varieties where every individual must be assigned to some
one of them, is a matter of some difficulty owing to the presence of
intergrading individuals. The characters which signalize var.
hispida Lauterborn and var. irregularis Lauterborn are not quickly
recognized under the conditions of rapid plankton enumeration, and
no effort was made to trace their seasonal distribution in our plank-
ton. Lauterborn's var. macracantha was included with the type
form his var. typica in our records. These two include those
individuals with medium-sized and longer posterior spines. In our
waters the variety macracantha is relatively rare, at least as figured
by Lauterborn ('98). Indeed, both the type and this variety consti-
152
tute less than a seventh of the total representatives of the species.
Their distribution throughout the year (Table I.) accords with the
results obtained by Lauterborn ('98), who found that the average
length of the posterior spine from January to May and from October
to December was from 78 to 48/i, while from June to September it
was from 28.5 to 21 //. In Table I. it will be seen that the longer-
spined forms which I have referred to A. cochlearis var. macracantha
and var. typica occur in the plankton from January to May 3 1 , and
then disappear, returning again, in small numbers, October. 25. The
short-spined variety referred by me to A. cochlearis var. stipitata
and the spineless var. tecta are, on the other hand, continued during
the summer. The natural result would be that the average length
of the spines in the species as a whole would fall during the summer
months. It is apparent that this tendency on the part of A.
cochlearis to become shorter and smaller during the summer months
does not bear out the contention of Wesenberg-Lund ('98) that
winter individuals are smaller and summer ones larger among
perennial rotifers. He reports var. tecta as "die Hauptform des
Winters " in several Danish lakes, and the variety with a long
horn as a summer form, found in July-August.
Of these varieties, macracantha, typica, and stipitata intergrade
in our waters with numerous connecting links, while var. tecta is not
connected with the other forms by many individuals with inter-
mediate characters. Lauterborn ('98) also notes the greater inde-
pendence of this variety in the waters of the Rhine.
In Table I. the seasonal distribution of these three varieties, the
.long-spined (typica and macracantha}, the short-spined (stipitata),
and the spineless (tecta} are given separately. It will be noted that
the long-spined form has the distribution above mentioned, that
var. tecta runs throughout the whole year, and that var. stipitata is
absent in midwinter and is a common summer form. The relative
numbers of the varieties fluctuate in different years. For example,
var. tecta was relatively but one fourth as abundant in 1897 as in
1898. As shown in Table I., whenever coincidently present in the
plankton all the varieties respond to the causes which produce the
rhythm of occurrence, the rise, culmination, and decline of the pulses
being much alike in all of the varieties.
About three eighths of the females noted in 1898 were ovigerous,
carrying as a rule but a single egg. Instances of two eggs were
153
noted, but they are rare. The greatest proportion of egg-bearing
females appears during the rise of the pulse, as is seen in the follow-
ing table, which gives the data of the vernal pulse in 1898. From
ANUR^EA COCHLEARIS.
Date
No. of
ovigerous
females
Total
females
Total
eggs
Ratio of
eggs to
individuals
No. of
dead
April 12
800
2,200
800
1 2 75
April 19
6,400
15,200
8,800
1 1.73
400
April 26
45 000
137 800
65 000
1 2 12
3 200
May 3 .
536,000
1,022,400
552,200
1 1.85
9 600
May 10
489 600'
1 145 600
643 200
1 1 78
99 200
May 17....
110,400
434,800
160 000
1 2 71
100 000
May 24
6,000
21,200
7,200
1 2.94
1,800
May 31.
3 000
11 200 *
3,400
1 3 29
1 800
April 12 to the crest of the pulse on May 10 (not inclusive) the aver-
age ratio of eggs to individuals was 1 to 1.87. From the crest to thp
foot of the decline inclusive the ratio is 1 to 2.98. The number of
empty loricus is given below, and it will be noted that on the week
prior to the crest of the pulse there w r ere 107 living to one dead ; on
the crest itself, one to twelve ; while the week following the crest of
the pulse there was an empty lorica for every 4.3 living females.
Rapid multiplication thus attends the rise of the pulse and rapid
destruction its decline. Parasites were very rarely observed in this
species. The decline of a pulse is thus due to the cessation of
reproduction and a relatively heavy death rate.
Apstein ('96) finds that in Lake Plon Anuraa reaches its maxi-
mum in July and is at its minimum in April. It is everywhere
common in the German w r aters. A. tecta, on the other hand, was
found only in the smaller lakes and in great numbers, replacing
cochlearis in warmer months to some extent. Lauterborn ('98)
regards it as the most abundant rotifer in the Rhine. Our statistical
records do not show that this is the case in the Illinois, for it is here
154
surpassed by several other species. Zimmer ('99) finds that this
species is the most common winter rotifer in the plankton of the
Oder, with a maximum in the spring and a predominance of var.
tecta from July to September. Schorler ('00) finds it to be the most
common rotifer in the Elbe from April to November ; and Skor-
ikow ('97) finds it in the Udy, in Russia, throughout the summer
in great numbers, but surpassed by Synchczta, Polyarthra, and
Brachionus angularis. The variety tecta greatly exceeds var.
stipitata in these waters. Seligo ('00) finds it throughout the year
in Prussian lakes near Danzig, with a maximum in May. There are
indications, in his data, of recurrent pulses during the summer, but
his interval of collection is too great to follow their history. Burck-
hardt ('OOa) finds it throughout the year in Swiss waters, with its
single maximum in August. Jennings ('94, '96, and '00) reports it
in the summer plankton of Lake Michigan and Lake Erie and of
inland waters of Michigan.
Anuraza hypelasma Gosse. Average number of females, 2,390;
of eggs, 1,917. This species has a very definite limitation to a
period extending from early in June to the first days of November.
There are but two records outside of these limits a single female and
egg on Jan. 11, 1898, and another upon April 19 of the same year.
The probabilities of occurrence in very small numbers at all tempera-
tures is thus indicated. The following table gives the data of pulses
and temperatures.
All of the pulses save one occur at temperatures above 70, and
with this exception the species declines rapidly and disappears
shortly after temperatures pass below 60. It is plainly, in our
waters, a summer planktont, with its optimum temperature close
to the summer maximum. This species takes no share in the vernal
pulse, and there is no satisfactory evidence of any fluctuation
corresponding to it at any other season. There are three or four
pulses in each summer, and the species is apparently poly cyclic, for
winter eggs were found in 1898 either at the maximum of the pulse
or the week or fortnight following. Thus 24,000 winter eggs were
recorded on Sept. 27, 1898, the date of the maximum of the Septem-
ber pulse. The parthenogenetic eggs preponderate during the rise
of the pulses in a very marked manner in this species. For example,
in this September pulse 55,400 eggs were recorded during its rise
to 500 during its decline. . In like manner, in the case of the
155
PULSES OF ANUR/EA HYPELASMA.
VpvQf
First
record
I
Dulses
Date
Temp.
Date
Temp.
No.
1896
June 2 7
80
June 2 7
80
1 ,200
1897
June 28
75
July 14
79
10,400
1898
June 14
83
June 2~1
77
9,600
Year
Pulses
Last record
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
1896
Aug. 15
" 29
81
74
2,000
3,600
Sept. 30
58
1897
Aug. 31
80
20,000
Oct. 5
71
23,200
Nov. 2
55
1898
Aug. 16
77
16,000
Sept. 27
Oct. 18
73
52
43 , 200.
13,500
Nov. 1
45
August pulse 15,200 eggs were found on the rise to 4,000 on the
decline.
The location of the pulses of A. hypelasma is of special interest.
It will be seen in Table I. that they occur in 1898 in the same col-
lections in which the pulses of the other species of Anuraa and many
other rotifers occur, or in collections but a week removed. They
coincide in general with dates of the ploiman maxima noted in the
opening discussion, and exhibit the same correlation with hydro-
graphic conditions and intercalation with the pulses of chlorophyll-
bearing organisms which were noted in the general discussion and
have been found in preceding species. The comparison with Anur&a
of the cochlearis group affords a curious instance of an entire sup-
pression (Table I.) of one species of a genus (cochlearis} in the month
of August and the occurrence of a normal pulse in another (hypelas-
ma}. Comparison of the distribution of cochlearis in previous
summers would lead us to expect a cochlearis pulse in August, 1898,
156
but none appears in this interval, while hypelasma runs a normal
course of recurrent pulses throughout the summer. This August
pulse of hypelasma (Table I.) culminates August 16, just a week
after the symmetrical and well-defined pulse of chlorophyll-bearing
organisms (PI. II.) of August 9.
With a single exception, all of the pulses of 1896 and 1897, indi-
cated in the table, fall a week later than, or coincide with, the pulses
of chlorophyll-bearing organisms, as in 1898 t
This species has not occupied a prominent place in the literature
of fresh-water plankton. Weber ('98) finds it rare in Swiss waters
in the summer. Lauterborn ('93) classes it with the monocyclic
summer forms in the plankton of the Rhine, though he states in a
footnote that he had" found winter eggs once in June. It is probably
poly cyclic in our waters. Skorikow ('96) finds it in the summer
plankton of the river Udy, in Russia, but it is not mentioned by other
investigators of the potamoplankton of Europe. Apstein ('96) does
not report it from Lake Plon.
Asplanchna brightwellii Gosse. Average number, of adults 2,079,
of eggs, 396; averages in 1897, 16,161 and 2,156. This is a poly-
cyclic perennial planktont in our waters. It has been found in
every month of the year, but the greater numbers and more con-
tinuous occurrences lie between May 1 and October 30. In 1898
(Table I.) all but 200 of the 108,120 recorded, lie within these limits,
and all but 260 above 60. In previous years approximately the
same limits are found. The following table gives the data of pulses
and temperatures.
PULSES OF ASPLANCHNA BRIGHTWELLII.
Year
Date
Temp.
No.
Date
Temp.
No.
1894
1895
June 19
June 27
80
80
6,678
1,600
1896
May 1
70
1,788
1897
1898
May 5
60
20,800
June 21
77
1,100
157
PULSES OF ASPLANCHNA. BRIGHTWELLII Continued.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1894
July 30
82
19,398
1895
July 29
75
1,344
Aug. 12
79
118,206
Nov. 14
45
1,725
1896
Aug. 2 1
79
1,200
1897
July 21
83
3,200
Aug. 10
81
5,200
Sept.- 9
. 80
284,000
1898
Aug. 2
79
23,200
Aug. 23
81
4,000
Sept. 27
73
6,400
It will be seen from this table that all the pulses save one, and
that one (Nov. 14, 1895) poorly denned, lie between 60 and the
maximum temperatures, indicating an optimum near the summer
maximum. There is in this species no prominent vernal pulse such
as that found in Anurcea, and the highest numbers were reached
during the height of the warm season.
The evidence of the polycyclic character of the seasonal distribu-
tion of this species is shown in the following table, which gives
the occurrences of ovigerous females, males, and winter eggs in 1898.
It will be noted that ovigerous females are more numerous during the
rise of the pulse; that the males appear just before, during, and
after the culmination of the pulse ; and that winter eggs are absent
only during the rise of the pulse, and appear at or after its culmina-
tion and during the decline. The data given afford a fine illustration
of the seasonal distribution of polycyclic rotifers, and of the relation
of the sexual cycle to the number and character of the representa-
tives of the species in the plankton. The growth of the pulse .results
from a rapid succession of parthenogenetic generations in the course
of about two weeks, and it culminates with or shortly after a pulse
in the food supply. The decrease in food supply is attended by the
appearance of males and winter eggs, a decrease in ovigerous
females, and a decline of the species. With the recurrence of the
food supply the parthenogenetic cycle again begins. The same
course of events is run in each recurrent pulse. Food supply
rather than temperature seems to be the determining factor in this
rhythm.
158
ASPLANCHNA BRIGHTWELLII.
Date
Males
Females
without eggs
Ovigerous
females
Winter
eggs
May 3
3,200
12,800
" 10
8,000
4,800
8,000
1 600
" 17
1 600
5 600
4 000
100
"24
400
200
" 31
200
400
June 7 ...
200
14
21
800
300
28
100
Tulv 5
120
40
120
12
o
o
19
40
240
" 26
240
12 400
5 260
60
August 2
4 000
7 200
12 000
5 600
9
80
800
16
800
800
23
3 200
800
60
30. . . ...
1 600
800
1 600
September 6
o
13
o
20
540
600
27
3 200
3 200
October 4
500
1 000
11
1 000
500
159
An examination of the location of the pulses of Asplanchna
brightwellii shows (Table I.) that in 1898 one coincided with the
pulse of chlorophyll-bearing organisms (PI. II.) and the remaining
four followed it either in a week or fortnight. In previous years
two pulses coincide with and five follow those of chlorophyll-bearing
organisms, and a single ill-defined one (Nov. 14, 1895) precedes.
This species is not wholly herbivorous in its feeding habits.
Codonella, Difflugia, and even other rotifers such as Brachionus and
Anur i -i M
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169
170
The reducing effect of the recurrent floods of 1896 may be traced
in the smaller numbers recorded in this year; and the larger num-
bers of 1897 may be referred to the more stable conditions then
prevailing. The very small numbers of 1898 may also be due to
disturbed hydrographic conditions of that year. The number is
much smaller than in 1896, when the hydrograph was even more
disturbed, but in this latter year there was more run-off of
impounded backwaters during the occurrence of B. bakeri,axuA this
would tend to favor their appearance in channel waters.
The occurrences and numbers of this species (as a whole) are
everywhere somewhat irregular, so that pulses of occurrence are
somewhat ill defined. Several such pulses are indicated in 1898,
and others recur in the records of previous years. As suggested by
the data of 1898 (Table I.), the several varieties share in these pulses.
The evidence upon this point is much more striking in other years,
when numbers are larger. For example, in the following table note
the pulse of 26,800 on August 23, 1897.
o
^j
to
Date
I
to
.S
^
to
Total
to
o
s
to
to
^>
S
8
p
S
fc
jt
E
to
. -o
O
to
**
-o
*
Aug. 10
o
o
200
200
" 18
1,200
200
600
2 000
5 200
7 400
" 23
o
7 800
1 800
3 400
2 200
11 600
26 800
" 28
200
400
200
1 000
1 800
In their location these pulses exhibit as a rule the same relation of
coincidence or sequence to the pulses of chlorophyll-bearing organ-
isms noted in some other species, and they frequently coincide with
those of other Ploima, but not always.
This is perhaps the most variable of the rotifers of the plankton.
At least its variations affect the fixed processes of the lorica and are
thus quickly and easily appreciated. The species, in common with
171
B. pala, B. angularis, and probably B. urceolaris, has a variety in
fact, several varieties with two posterior spines which are usually
symmetrically placed but not always symmetrically developed. The
form without posterior spines (var. dumorbicularis Skorikow) inter-
grades with these, and a series might be formed with complete
intergradations linking this in turn with var. rhenanus Lauterborn,
in which the spines are but slightly and often unequally developed.
From this we pass, by a slight elongation of the posterior spines, to
var. brevispinus Ehrbg., thence to the type in which- the spines as
figured by Rousselet ('97) are directed .posteriorly with but slight
curvature. From this we may pass toward variants in which the
symmetry is preserved, but the spines are much elongated and
curved outwardly. The anterior spines in such individuals are also
more elongated and exhibit a similar outward curvature (var.
melhemi Barrois and v. Daday). Extreme types of this curvature
sometimes occur (J5. falcatus Zach.). In another direction we find
the bilateral symmetry of the processes, both anterior and posterior,
to some extent lost as a result of differences in the curvature of the
spines (var. tuberculus Turner). There are also differences in the
surface markings of the lorica which have been utilized as specific
distinctions. Kertesz ('94) describes as B. granulatus a species
with a minutely pustulate surface, and Turner's B. tuberculus takes
its name from this same feature. It seems questionable, however,
if these surface markings are even of varietal value. Individuals
without spines, in which the transverse diameter is relatively large
(var. obesus Barrois and v. Daday) , are also found.
In assorting the individuals belonging to this variable group I
have arranged them under the following heads: bakeri O. F. Mull.,
bidentata Anderson (non bidentatus Kertesz), brevispinus Ehrbg.,
cluniorbicularis Skor., melhemi Barrois and v. Daday, obesus Barrois
and v. Daday, rhenanus Lauterborn, and tuberculus Turner. The
number might have been increased. The individuals referred to
var. melhemi include many if not all of the long-spined specimens
such as Rousselet ('97) has referred to the type, the latter designa-
tion having been given to individuals intermediate between this and
brevispinus. The variety tuberculus includes the asymmetrical
individuals, regardless of the surface markings. I will now briefly
compare the seasonal distribution of these varieties and note
any peculiarities which mark them individually :
172
Brachionus bakeri O. F. Mull., type form. Average number, 2.
As shown in table on p. 1 93 (MS.), this form is much more abundant
in previous years though it is relatively rare, ranking sixth in the list
of seven forms recognized. The most of the records fall prior to the
middle of August, and it seems to be an early rather than a late
summer form.
Brachionus bakeri var. obesus Barrois and v. Daday. Average
number of females, 41 ; of eggs, 62. The proportion of egg-bearing
to non-egg-bearing females 2 to 3 in all records is larger than in
any other variety. It seems probable that the lateral expansion
which marks this variety may be only the result of rapid reproduc-
tion. In common with most of the other varieties this one occurs
at the time of the pulses, but it is last in the list of seven, and the
numbers are too small to trace its seasonal preferences with cer-
tainty.
Brachionus bakeri var. bidentatus Anderson (non Kertesz).
Found once August 5, 1895, at 78.
Brachionus bakeri var. cluniorbicularis Skor. Average number
of females, 90; of eggs, 95. This also was more abundant in all
previous years. This variety is, next to tuber culus, the most
abundant of the varieties in our plankton. The two stand at
opposite extremes of the series of varieties, the former being least
modified, and the latter most, especially in the direction of asym-
metry. It includes about one third of all the individuals of the
species. The ratio in the grand total of females to eggs carried
11,708 to 5,976 is somewhat less than the average in the entire
species. This variety is distributed throughout the whole seasonal
range of the species with no marked predominance in any particular
part of it. It is wholly absent in the early summer of 1 897 , but very
abundant in late summer of that year, though not in other years.
The autumn of 1897 was one of long-continued high temperatures
(Pt. I., PL XL), and under those conditions this variety constituted
two thirds of the individuals belonging to the species. If we add to
it the representatives of rhenanus, obesus, and brevispinus we have
a total of 15,400 individuals with no posterior spines, or with spines
but slightly developed, in contrast with only 2,200 with such well-
developed spines referred to varieties melhemi and tuberculus. The
conditions of temperature were those in which according to the
173
hypothesis of Wesenberg-Lund ('00) we should expect a predomi-
nance of the long-spined forms.
Brachionus bakeri var. rhenanus Laut. Average number of
females, 118; of eggs, 138; but more abundant in previous years.
This is the third in numbers on the list of seven varieties, being
surpassed only by duniorbicularis and tuberculus. It includes about
one sixth of the individuals referred to this species. ' It is found
throughout the whole range of the seasonal distribution of the
species and exhibits the same peculiarities noted in-cluniorbicularis,
to which it is very closely related. The proportion of females to
eggs noted in this variety is very large; 5,284 to 5,485 in the grand
total.
Brachionus bakeri var. brevispinus Ehrbg. Average number of
females, 795 ; of eggs, 390; but somewhat more abundant in previ-
ous years. It was found throughout the whole seasonal range of
the species, but not quite so abundantly in the latter as in the earlier
half of the summer, resembling in this particular the type. The
number of eggs carried in this species is in relation to the number of
females less than usual 3,906 to 795.
Brachionus bakeri var. melhemi Barrois and v. Daday. Average
number of females, 49; of eggs, 49. More abundant in previous
years, especially in 1894, when it constituted over a fifth of the
individuals (25,764) in the largest pulse recorded for the species as a
whole 122,958 on July 30. In the aggregate in all years it includes
only about a ninth of the individuals referred to the species. This
form was originally described from Syria, but it is found in great
perfection in our plankton, even in the extreme type described by
Zacharias ('98b) as B. falcatus. It occurs throughout the whole
seasonal range of the species, its distribution being somewhat similar
to that of tuberculus. I do not find any constant tendency limiting
its occurrence to any part of the seasonal range.
Brachionus bakeri var. tuberculus Turner. Average number of
females, 155 ; of eggs, 42 ; but very much more abundant in previous
years, especially in 1894, when it constituted almost half (55,332) of
the largest pulse of the species (122,958). This, the most divergent
of all the varieties, constitutes over a third of all the individuals
referred to the species. It occurs throughout the whole seasonal
range of the species, though the larger numbers were found in
1894-97 in the earlier part or middle of the summer. I find nothing
174
in a comparison of the seasonal distribution of these more decidedly
spinous varieties of B. bakeri with that of the smoother forms, such
as cluniorbicularis , which indicates any correlation with temperature
conditions of a nature to support Wesenberg-Lund's suggestion
that the elongation of the processes of plankton organisms arises in
response to the lessened buoyancy of the water during higher tem-
peratures. Forms with and without such processes are found among
the varieties of this species, and both occur indiscriminately through-
out the whole range of seasonal occurrence, and, so far as I can see,
the statistical data of their distribution with respect to temperature
afford no evidence of a correlation of spinosity and high tempera-
tures in this species. Other factors doubtless enter into this
problem and obscure this response if it exists.
B. bakeri is everywhere widely distributed in fresh water. Its
occurrence in the plankton of open waters has not, however, been a
matter of frequent note. In fact there is some reason to think that
it is largely confined to shallow warm waters where vegetation is
close at hand, or where at least the flagellates and smaller algae
abound, as they do in water fertilized by decaying vegetation or
other organic matter. There is, it seems, no reason for regarding
this species as merely adventitious in our plankton. It bears all the
characteristics of a true limnetic organism in our environment. Its
presence in the plankton is not due to floods or other disturbances
which might carry it from a littoral region into the open water. It
exhibits characteristic pulses, and is found everywhere in summer
in company with typical planktonts in open water.
Zacharias ('98) records it in some German ponds and streams,
and Weber ('98) in Swiss marshes in the warmer months. Stenroos
('98) also finds it in the summer plankton of littoral and open waters
in the shallow Nurmijarvi Lake in Finland. Jennings ('00) reports
it as one of the commonest rotifers in East Harbor, Lake Erie, and
in the swamps on the islands. In land-locked pools short-spined
varieties were found, and in swamps the long-spined. Speaking of
this difference, Jennings says " Possibly the different form found in
these pools is due to the greater concentration of various salts in this
water or to some kindred factor." In our own region both varieties
occur at the same time in the same environments, channel and
backwaters alike, and such factors as Jennings suggests to explain
the appearance of the varieties cannot well be operative here in
175
channel waters. Schorler ('00) reports the species as sporadic in
the Elbe, and Skorikow ('97) finds both B. bakeri and its variety
brevispinus sparingly in the Udy in summer months.
This species in common with other Brachionida: was infested by
Bimcerium hyalinum Przesm., and occasionally by a filamentous
fungus-like growth. Empty loricae were wont to appear with the
culmination of a pulse and subsequently. No males were identified
as belonging to this species, and attached male eggs were recorded
only late in September, 1897, at the close of an unusual pulse. They
were found on var. cluniorbicularis and rhenanus. Females with
winter eggs were not at any time recorded for this species. It may
be that some of the free winter eggs referred to the genus Brachionus
(Table I.) belong to this species. The recurrent pulses are similar to
those of known poly cyclic species, and we may infer the probability
of such a phenomenon in B. bakeri, though conclusive proof of its
occurrence is not found in the statistical records.
Brachionus budapestinensis v. Daday. Average number of
females, 4,211; of eggs (carried), 740. This is one of the most
sharply defined species of Brachionus and a typical planktont of
open waters. It has, moreover, a sharply limited seasonal distribu-
tion in which it is apparently poly cyclic. The appended table gives
the dates and temperatures of appearance and disappearance and
the pulses in the several years.
In the main, the period of occurrence is practically from the end
of June till the early part of October and above 60. A record in
May, 1896, and an isolated one in December of the same year, indicate
an extension of this period, but such occurrences are rare and
irregular and the numbers small. This abrupt decline in 1898 as
temperatures pass 60 (PI. XII., Pt. I., and Table I.) is paralleled
in previous years. The normal seasonal routine seems to be as
follows: The species reappears in the plankton in May- June at
70, rising slowly to its first pulse (average, 26,104) in July, with a
larger pulse (average, 184,453) in the following month during the
maximum heat, and a much smaller one (average, 10,044) in Sep-
tember, followed immediately by an abrupt decline. The average
temperature of the larger pulses lies close to the season's maximum,
while the latest pulse, at the lower temperature (72.2) averages but
10,044. These data all indicate that this is a midsummer planktont,
with its optimum temperature near the summer's maximum. The
1
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1
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176
177
relation of hydrographic conditions to the relative development of
pulses in different years is seen on a comparison of the record for
1896 and 1897, the former (Pt. I., PI. X.) being a year of recurrent
floods and the latter (Pt. I., PI. XI.) one of stable conditions
through the greater part of the seasonal distribution of the species
in question. The average numbers in these two years were 3,105
and 31,306, respectively, and the average amplitude of the pulses
18,250 and 97,200, showing, respectively, a ten- or five-fold increase
in the latter year. The extension of the heated termjnto September
in 1897, is reflected in the large September pulse (552,000) and in
the extension of the period of occurrence into October.
The locations of the pulses of Brachionus budapestinensis in 1898
correspond with those of the Ploima in general. They likewise
coincide with or follow those of the chlorophyll-bearing organisms
(cf. PI. I. and II. with III. and IV. and Table I.). Similar relations
are apparent in 1896 and 1897 but are less evident in prior years.
They suggest an interrelationship of the pulses in this species with
the fluctuations in the food supply.
Males, male eggs, and winter eggs were not recorded, but the
recurrent pulses in this species are so similar to those in other rotifers
in which the evidence of the occurrence of sexual reproduction at
the culmination of each pulse has been found, that the inference
may be made that this species likewise is poly cyclic in our waters.
Females carrying one or two summer eggs have been found in
greatest abundance during the rise of the pulse, and only in small
numbers, if at all, during its decline.
This species is subject to some variation in the development of
surface ornamentation, in the ratio of width and length, and in the
curvature of the median spines. It is usually somewhat more
slender than figured originally by v. Daday ('85) or even by Hempel
('96) , who described a form somewhat more slender than that figured
by v. Daday, as B. punctatus. Shortly afterwards Skorikow ('96)
described the same species as B. lineatus from Russian waters. The
name given by v. Daday has, priority, and as neither the Russian nor
the American forms are to my mind well enough set off to merit
even varietal distinction, I have used the name given by v. Daday,
and have included under it both wide and narrow forms and those
with incurved or outcurved median spines. The fact that their
common record of seasonal distribution forms a seasonal curve of
178
typical character is corroborative of the view, though not conclusive,
that we are dealing with a single species and not with several.
This species has not been widely reported in the fresh-water
plankton. It is evidently a planktont of warmer waters, and for
that reason may have escaped notice, since the cooler waters have
been the more thoroughly explored. Thus it was not found by
Weber ('98) in Swiss waters in his thorough explorations about
Geneva, nor by Jennings ('94, '96, '00) in the Great Lakes or inland
waters of Michigan. It has, however, been recorded by Skorikow
('97) in the plankton of the Udy River, in Russia, where it was
exceeded in number by only two species of its genus, B. pala and
B. angularis, ranking tenth in numbers among all the rotifers. His
data of frequency from July to October suggest several recurrent
pulses. It has likewise been found by Lauterborn ('98) in the
plankton of the Rhine, where he classes it with the stenothermal
planktonts. Zacharias ('98) finds it in ponds near Leipzig, and it
was originally described by v. Daday ('85) from Hungarian waters,
and again noted there by Kertesz ('94). Fuller exploration of the
summer plankton in warmer regions will doubtless extend the record
of its range.
Brachionus militaris Ehrbg. Average number of females, 147 ;
of eggs (carried), 98. In previous years the species was much more
abundant, the averages in 1897 being 1,412 females and 523 eggs, and
in 1896, 1,288 females and 576 eggs. This greater development in
years prior to 1898 is evident in many of the Brachionidcz.
The following table gives the dates of first and last records in
each season, and the location, temperature, and amplitude of the
pulses in the several years.
This is evidently a summer planktont with well-defined limits.
These limits appear much less evident in 1898 (Table I.) than in
prior years. In 1896 and 1897, for example, the species is almost
continuously present in the plankton from the time of its first
appearance until the last record for the season. All of the records
save two lie above 70, and the average temperatures at which the
pulses occur are all at or above 80. Its optimum thus lies near
the summer maximum. The lower limits are not definitely
established owing to insufficient collections in periods of rise
and decline, but they seem to lie near 70, with small numbers
lingering to 60.
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180
This species has never developed large pulses in the channel
waters of the Illinois. Hempel's statement ('99) that it is "the
most abundant species of the genus" can apply only to certain
collections in vegetation-rich backwaters, for in the river it is sur-
passed in the totals of occurrences in the statistical records by eight
other forms of firachionus, namely, variabilis, pala, amphiceros,
dorcas, rubens, budapestinensis, duniorbicularis, and tuberculus. I
found it in very great abundance in the July-August plankton of
Crystal Lake, a shallow warm pond rich in vegetation, formed by
damming a small creek tributary ta the Wabash system, near
Urbana, 111. From the relatively small numbers, the slight ampli-
tude of the pulses, and their somewhat irregular development I am
inclined to think that the centers of distribution of this species are
not in the open water of the river and its backwaters, but more
in the vegetation of warm, shallow regions such as the margins of our
bottom-land lakes. It is thus to some extent adventitious in our
plankton.
The pulses of this species are relatively so small that they do not
contribute an appreciable amount to the total ploiman pulses, nor
do more than 50 per cent, of their number coincide with such general
pulses, though they are sometimes found during their rise. The
greater part of them coincide with the pulses of chlorophyll-bearing
organisms (PL I. and II.), suggesting a food relationship.
This species is one of the best-defined in the genus, though in the
character of its asymmetry it varies toward B. bakeri var. tuberculus
Turner. It exhibits some variation in the degree of asymmetry, in
the curvature of the spines, and in the surface markings. The indi-
cations of pulses suggest a poly cyclic habit, but no evidence in the
way of males, male eggs, or winter eggs was recorded which will
substantiate the inference. A female carrying a winter egg w r as
found Sept. 21, 1-897, at the close of the period of occurrence. Fe-
males with one, two, or three summer eggs were found throughout
the summer and in somewhat larger numbers during the rise of the
pulses.
Brachionus mollis Hempel. Average number of females, 137;
of eggs, 10. More abundant in previous years, the average in 1897
being 1,092 and 277, and in 1896, 428 and 56.
This likewise is a summer planktont. The earliest record of its
appearance in the plankton is June 17, 1896, at 76; and the latest,
181
October 17, 1894, at 58. With but two exceptions the species was
taken only above 70, and the period of most continuous occurrence
and largest numbers is near the summer maximum of 80. The
optimum is thus near the summer maximum. This species was
never taken in the plankton in large numbers, the greatest being on
Sept. 14, 1897 (20,000), at 84. On account of the small numbers
.and somewhat irregular occurrences the phenomenon of recurrent
pulses is here less apparent than it is in more abundant species. The
appended table records the best-defined ones. Thesejpulses share in
the general ploiman pulses in only about 50 per cent, of the cases,
and the most of them coincide with or follow shortly after the pulses
of chlorophyll-bearing organisms.
PULSES OF BRACHIONUS MOLLIS
Year
Date
Temp.
No.
Date Temp.
No.
1895
July 6
81
742
Sept. 5 75
954
1896
July 18
79
1,200
Aug. 21 \ 79
8 , 400
1897
July 30
85
1 1 , 600
Sept. 7 80
20,000
1898
Aug. 23
81
800
Sept. 27 73
4,800
So far as I am aware this species has not been found in other
waters than the Illinois River and its adjacent backwaters. Hempel
('99) reports it as most abundant in the marshy environment of
Flag Lake.
Brachionus pala Ehrbg. Average number, including all varie-
ties: females, 19,969; eggs, 25,974. The following table gives the
average number, in the several years, of the varieties here included,
and it will serve to show their relative frequency.
This is the most abundant species of the genus in our waters, the
grand total of all occurrences exceeding 9,000,000. As a whole the
species was much more abundant in the stable year 1897 (180,998),
and less abundant, all things considered, in the disturbed conditions
of 1896 (36,665). As a whole the type form pala is less abundant
than amphiceros. It forms but 28 per cent, of the total, as compared
with 68 per cent, included in the latter variety. Dorcas forms less
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culmination of a pulse is very large. For example, the sudden ver-
nal pulse of 716,982 on April 24 is accompanied by 28,584 free
winter eggs. The pulse declines to 22,224 on April 29, and the free
winter eggs rise to 95,841, and the empty loricas to 26,114.
Females carry 1-5 summer eggs, and 1-8, or even more, male
eggs. There is great variation in the size of the summer eggs, these
and -the male eggs appearing almost to intergrade.
Brachionus pala, including B. amphiceros, is a common constitu-
ent of the plankton of shallow warm waters. It has not been
reported from the larger and cooler lake waters by Apstein ('96),
Burckhardt ('00 and 'OOa), or Jennings ('94, '96, and '00). Zacha-
rias ('98) and Marsson ('00) find it in the summer plankton of
smaller lakes and ponds in Germany. Seligo ( '00) records it from
April to October, with a maximum in August, in Prussian lakes ; and
Lauterborn ( '98a) finds it to be perennial and polycyclic in -the
Rhine. Schorler ('00) reports both pala and amphiceros from the
Elbe, the former being abundant in May and sporadic during the
summer, while the latter was abundant in April, June, and Septem-
ber, and rare at other times during the warmer months. Zimmer
189
('99) finds amphiceros in the Oder, where it appears in April and.
increases until the end of August or the first of September, when
it is the most abundant animal in the plankton. In no one of these
instances was the examination so long continued or made at such
short intervals as in the case of the exploration of the Illinois. The
diversity exhibited in these different waters may be paralleled by
the fluctuations from year to year in the Illinois, and from all the
data it may be inferred that the organism is probably perennial and
polycyclic, the number of pulses depending upon local conditions,
primarily of the food supply.
Brachionus pala var. amphiceros Ehrbg. Average number of
females, 17,071; of eggs, 5,103. The numbers were much larger
(158,299 and 35,392) in the stable conditions of 1897, and still
smaller (5,430 and 715) in the disturbed conditions of 1896.
The seasonal distribution of this variety with respect to that of
the type constitutes the chief point of interest in the records. It is
present throughout the whole range of temperatures, shares in the
vernal and autumnal pulses noted for the species as a whole, but
constitutes a much greater proportion of the amphiceros-pala group
during the warmer months than it does in the colder ones. Thus,
as shown in the accompanying table, the proportion which amphi-
SEASONAL DISTRIBUTION OF BRACHIONUS PALA AND B. PALA VAR. AMPHICEROS.
Year
June 1 to Oct. 1
Oct. 1 to June 1
pala
amphiceros
pala
amphiceros
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196
197
ber 1, but is continuously present in the winter of 1898-99 from
December 6 till March 28, when collections ceased.
Male eggs were recorded but once April, 29, 1895 and there
is no other evidence of the cycles of reproduction beyond the pulses
in numbers. They suggest a polycyclic habit with major pulses
in spring and fall. It is apparent that conditions affect these cycles
greatly, as is seen, for example, in the contrast between the earlier
years, with low water in the spring, and the later ones, when high
water was longer continued.
This variety, rubens, has not been widely reported in the plank-
ton. Skorikow ( '96) finds it in June in the River Udy, and Kertesz
( '94) reports it from Budapest, while Stenroos ( '98) finds it in the
littoral fauna of Lake Nurmijarvi in Finland, and also in the plank-
ton in July and August.
Brackionus urceolaris var. bursarius Barrois and v. Daday.
Average number of individuals, 206; of eggs, 33. This is a sum-
mer variety, and forms but a small part less than one per cent.
of the total number of individuals referred to the species.
Brachionus variabilis Hempel. This species was found but once
in 1898, but was more abundant in former years (see table on oppo-
site page). The largest development which it attained in the Illinois
was a pulse of 168,222 on August 15, 1894, at 84. The largest
number in subsequent years was 5,200 per m. 3 on August 8, 1896.
It may be significant of the connection of this form with the urceo-
laris-rubens group that the great pulse of 1894 was coincident with
an unusual development of rubens on that date.
This species is a summer form, the earliest record being May 24,
1898, at 74, and the latest September 25, 1895, at 73. Its opti-
mum temperatures lie near the summer maximum. If this form
should prove to be merely a spinous variety of B. urceolaris it will
afford another illustration of spinous varieties of Brachionus appear-
ing at high temperatures, in accordance with the hypothesis of
Wesenberg-Lund ('00).
In Table I. there is given for 1898 the seasonal distribution of
the free winter eggs of Brachionus. It will be seen that they occur
throughout practically the whole year, with some increase after the
times of the April-May and September pulses.
Cathy pna leontina Turner. Average number, 47, in 1896, a year
of disturbed hydrograph ; less abundant in previous years, and not
198
recorded in subsequent ones. Earliest record, June 17, at 76; and
latest, October 2, at 63. Always present in small numbers and
evidently adventitious.
Cathy pna luna (Ehrbg.) Gosse. Average number, 47. Found
in every month but November, though always in small numbers and
irregularly. All but six of the thirty-three records fall between
April 1 and October 3 and above 50. Over half of all the individu-
als were found in 1896. This fact, together with the nature of the
seasonal distribution, indicates plainly its adventitious character.
Cathypna rusticula Gosse. Found once, March 22, 1897, at 44.
Not previously reported from American waters.
Ccelopus porcellus Gosse. Average number, 106. From March
to September, at 37 to 80, and apparently adventitious.
Colurus bicuspidatus Ehrbg. Average number, 274. This
species is apparently a winter planktont. In 1897 it appeared first
November 9, at 50, and was found somewhat irregularly through
the winter until May 17, at 64. There is a pulse March 15, at
46, of 6,400. Ovigerous females were found during the rise of the
pulse, and males on April 12, on its decline. A few scattered records
were made in the following winter, beginning November 8, at 46. It
occurs in the plankton during flood season and may be adventitious.
Colurus obtusus Gosse. Average number, 38. In small numbers
and irregularly in March and April at temperatures below 50, and
in September at 73. Hempel ('99) lists also C. deflexus Ehrbg.
Diglena circinator Gosse. Average number, 121, in 1896, a year
when many adventitious rotifers were brought into the plankton
by disturbed hydrographic conditions. All the records lie between
April 29, at 70, and July 28, at 81. An ovigerous female was found
in July. The species is adventitious in the plankton.
Diglena forcipata Ehrbg. was recorded once October 12, 1897,
at 65.
Diglena giraffa Gosse was observed but once in the river plank-
ton. ^Not before recorded from American waters.
Diglena grandis Ehrbg. was recorded in July and September at
76 and 79.
Diglena uncinata Milne was found August 12, 1898, at 82.
Hempel ('99) reports D. biraphis Gosse and D. catellina Ehrbg.
in waters immediately tributary to the river. All members of the
199
genus belong to the littoral fauna among vegetation, and are adven-
titious in the plankton of open water.
Euchlanis pyriformis Gosse. Recorded April 12, 1898, at 52.
Hempel ('99) reports it from June to October in collections in the
river in 1894 and 1895.
Euchlanis triquetra Ehrbg. Average number, 19. Found irregu-
larly from July to November at 84 to 41. Hempel ( '99) reports it
also in June. It is probably adventitious.
Hempel ('99) also reports E. dilatata Ehrbg. in, the river from
July to September, and E. deflexa Gosse in tributary waters.
Gastropus stylifer Imhof. A rotifer doubtfully referred to this
species was found sporadically in the plankton of the river. It was
recorded in June, 1894, and July, 1896, at temperatures above 75.
It was almost continuously present in 1896 from February 20 to
April 10, and again on November 17 and December 3. It did not
reappear until January 31, 1899, from which time it continued
present until the close of operations in March. Most of these oc-
currences are at minimum temperatures and all of them below 45.
I have followed Weber ('98) and Jennings ('00) in using Imhof 's
name Gastropus stylifer instead of Hudsonella picta Zach. or Notops
pygmcEUs Caiman, by which names the species has been frequently
designated. The evidence from our records indicates that it is a
somewhat sporadic winter planktont in our waters. Lauterborn
('93) finds it to be a perennial planktont in the Rhine, with its
largest numbers in summer.
Hydatina senta Ehrbg. was found September 20 at 73. Hempel
('99) also reports it in towings from the river in March and July,
1895. This species is very common in European waters, but has
as yet been found in America only in the Illinois River and, by Kel-
licott ('88), at Corunna, Mich.
Mastigocerca bicornis Ehrbg. Average number, 42. Found
irregularly and in small numbers from June 28 to September 13
above 63. Hempel ('99) reports it from Quiver Lake among
vegetation, and it is evidently adventitious in the river plankton.
Mastigocerca bicristata Gosse was found but once, late in Septem-
ber, 1895, at 73, but it is more abundant in the backwaters.
Mastigocerca carinata Ehrbg. Average number, 1,674. This
species was present in the plankton from the middle of June till the
200
first of October, and at irregular intervals and in small numbers in
fall and winter months. The distribution in years prior to 1898
falls within the limits shown in Table I. In this year the bulk of
the occurrences lie between June 21 and August 4, and above
77 and 72. The optimum lies near the summer maximum, though
occurrences at minimum temperatures in March and December
reveal acclimatization to a wide range of temperatures. In this
year there are several somewhat irregular pulses, the best-defined
of which follow the pulses of chlorophyll-bearing organisms (cf.
Table I. and PI. II.) at an interval of one or two weeks. The species
was not recorded so frequently in previous years, in some of which
also pulses are indicated. These pulses are not consequent upon
floods, and the species is apparently not adventitious in the plankton
but a normal constituent. Apstein ('96) reports M. capucina as
abundant in Dobersdorfer Lake from June to October a seasonal
distribution similar to that found in the Illinois River for M.
carinata.
Mastigocerca elongata Gosse was found once March 28, 1899, at
38. Hempel ('99) reports it in June in Quiver Lake.
Mastigocerca mucosa Stokes was taken in August to October,
1898, at 82- 62, in small numbers. It is reported by Jennings
('00) as "one of the most abundant of the Rotifera among the
vegetation of the shallow parts of Lake Erie," but it was not reported
by Hempel ('99) in similar environment about Havana.
Mastigocerca stylata Gosse was found -in the plankton in small
numbers in June and July at temperatures approaching 80. Hempel
('99) reports it also in August.
In addition to the species of this genus above listed, Hempel ('99)
records M. lata Jennings. There are also in our records a considera-
ble number of individuals referred to this genus but not specifically
identified. Many of these belong to one, or possibly several, very
small species. They are most abundant during the summer months,
reaching a pulse of 16,800 on June 28. They occur in large numbers
in the filter collections (average for 1898, 798 ; filter-paper, 145,384),
and, it seems, must escape with ease through the silk net on account
of their small size and their active movements.
A number of species in this genus have been described of late
from the fresh-water plankton, but in the present state of the litera-
ture of the subject I am not certain to what species these forms
201
should be referred. The genus is sadly in need of critical revision.
It includes a number of semi-limnetic species, whose importance in
the plankton will probably be revealed by more perfect methods of
collection.
Metopidia lepadella Ehrbg. was found only in March and June
at temperatures above 46. It is apparently adventitious.
Metopidia oblonga Ehrbg. was found once July 29, 1895, at 75.
Metopidia salpina Ehrbg. w r as recorded June 28, 1898, at 78.
Metopidia solidus Gosse. Average number, 67. This is the
most abundant representative of the genus in our plankton. It was
recorded from March 15 to November 14, at temperatures above
45. Most of the occurrences are in the summer months (Table I.),
at maximum temperatures. The numbers are small, the occurrences
irregular, and the species evidently adventitious.
M. rhomboides Gosse is recorded by Hempel ( '99) from the river
plankton, as also M. acuminata Ehrbg., triptera Ehrbg., and bractea
Ehrbg. from the backwaters.
Monostyla bulla Gosse. Average number, 50. Present in small
numbers and irregularly from April till the middle of October at
temperatures above 50. It is evidently adventitious. Jennings
( '00) finds. this one of the most abundant rotifers among the aquatic
vegetation in Lake Erie. It is in our waters the most abundant of
the genus in the plankton, especially in the vegetation-rich back-
waters.
Monostyla lunaris Ehrbg. Average number, 37. Found in the
extremes of the temperature range, but over 50 per cent, of the
occurrences are in August-October. Its numbers are always small
and its occurrences irregular. It is plainly adventitious.
Monostyla quadridentata Ehrbg. Average number, 10. This
species was found in the plankton irregularly in July-September, at
maximum temperatures. It is abundant (Hempel, '99) in the
backwaters, where vegetation is abundant, and is apparently adven-
titious in the plankton. In addition to the species here recorded
Hempel ('99) lists M. cornuta Ehrbg. and M. mollis Ehrbg. from
collections in the river, and M. dosterocerca Schmarda from the back-
waters. This is an exceedingly variable group, and will repay a
thorough revision in the light of a study of the variation of its
species. A considerable reduction in the number of these so-called
species will doubtless result from such a study.
202
Noteus quadricornis Ehrbg. Average number, 19. This is a
rare species in the plankton, being found in 1895 and 1896 in July at
maximum temperatures, and in 1898, on April 12, at 52, and on
November 8, at 46.
Notholca longispina Kell. This species, which has been found
in the summer plankton of many European and American waters,
especially our Great Lakes, was noted but once in the Illinois in
January, 1895 (Hempel, '99). It seems to prefer cooler and purer
waters.
Notholca striata Ehrbg. Average number, 437, including varie-
ties. This is a winter planktont in our waters, appearing in 1897 on
November 30, at 34, reaching a maximum of 10,840 March 22
(Table I.), at 51, and disappearing April 19; at 52. It reappears
the following autumn on November 1, at 45, and attains a maxi-
mum of 4,000 March 21, at 37. In previous years the occurrences
all lie within the limits of November 1 and April 24 with the excep-
tion of two records in 1895 September 5 and October 15, at 74 and
56. The spring maximum in 1896 (7,778) was on April 10, at
52, and in 1897 (4,260) on March 22, at 43. In each year but a
single pulse, that of March-April, is indicated. Minor fluctuations
during the winter (Table I.) are in some cases attributable to flood
agencies.
The temperature limits of this species are quite definitely estab-
lished. The species reappears in autumn when 45 is reached, and
declines rapidly in the spring after 50 is passed and is but rarely
found above 60. It attains its greatest numbers late in winter or
early in spring in the face of flood conditions, though the numbers
attained in the channel waters are never very large.
Empty loricas have been found in the plankton after the decline
of the species in April, and females with a single egg were noted in
small numbers in 1895 during the rise of the pulse.
I follow the suggestion of Weber ('98) that N. striata should
include as varieties the following: N. labis Gosse, N. jugosa Gosse,
and N. acuminata Gosse. Examination of many individuals in the
plankton proves beyond a doubt the great variability of the organ-
ism whose seasonal occurrence we have traced. It varies in the
length of the posterior spine, in the proportions of the lorica, and
in the development of the striae and the anterior spines. Of a total
of 81 ,227 of Notholca striata in this wider sense, 68,887 were referred
203
to var. acuminata, 3,852 to var. jugosa, 7,029 to N. striata in the
narrower sense, and 1,469 to other varieties, including var.labis and
var. scapha. The seasonal distribution of N '. striata (sensu strictu)
and var. jugosa lies within the limits of that of var. acuminata, but
occurrences are too few to trace their seasonal fluctuations.
This species is reported by Lauterborn ('94) in the winter
plankton of the Rhine. He also notes the connecting links between
N. acuminata, N. striata, and N. labis, and regards them as belonging
to the same " Formenkreis." Apstein ('96) reports N. acuminata,
N. labis, and N. striata in lakes of northern Germany and indicates a
seasonal distribution which coincides closely with that found for
these forms in the waters of the Illinois. He also reports a March-
April maximum and only isolated occurrences in midsummer.
Forbes ('83) finds the species in the stomachs of young Coregonus
feeding upon the March plankton of Lake Michigan. Seligo ('00)
also finds it in the winter plankton of Prussian waters.
Notommata cyrtopus Gosse was found in the plankton in April
and September at temperatures above 50. Hempel ('99) reports
A r . aurita Ehrbg. from the river, and N. tripus Ehrbg. and N. lacinu-
lata Ehrbg. ( = Diaschiza lacinulata Ehrbg.) from the backwaters.
Ploesoma lenticulare Herrick was found in the plankton of the
river from September to December, 1896, throughout the whole
range of temperatures from 75 to the winter minimum. Hempel
( '99) reports it from May to December, but principally in vegetation
Polyarthra platyptera Ehrbg. Average number of individuals,
86,674; of eggs, 52,560. In 1897, 94,653 and 58,235 ; in 1896, 29,653
and 11, 138; in 1895, 28,947 and 20,074; in 1894, 743 and 217. The
effect of the stable conditions of 1897 and of the recurrent floods of
1896 is seen in the larger averages in the former year and in the
smaller ones in the latter.
This is one of the most abundant rotifers in our plankton, includ-
ing, as it does, one seventh of the total Rotifer a, and exceeding in
numbers all other species of the group excepting only Synch&ta
stylata. It is a perennial form, and was recorded in every plankton
collection but two, and it may have been present then.
The seasonal distribution of this abundant species is very char-
acteristic of the form which most, though not all, plankton organ-
isms exhibit. Two prominent features are (1) a limitation of large
numbers to the warmer months and (2) a rhythmic occurrence of
204
recurrent pulses at approximately monthly intervals. In Plate V.
I have plotted the seasonal distribution of this species for the years
1894-99. The plate will serve as one of the best illustrations of
the nature of the data contained in my statistical records that could
be chosen from them. It illustrates graphically the character of
the seasonal distribution of this species and the nature of what I
have called recurrent pulses.
In the table which follows, as elsewhere in similar tables, these
pulses are listed by the number of individuals attained at their
maxima, and are located according to the dates of these maxima.
PULSES OF POLYARTHRA PLATYPTERA.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1896
Jan. 6
" 25
32
33
5,406
2,736
Feb. 25
34
7.852
Mar. 24
41
57,267
Tan 9?
Feb 22
39
1899
Jan. 17
33
20,800
Feb. 14
33
145,600
Mar. 7
33
71,200
Year
Date *
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1896
Apr. 24
72
233,436
May 8
76
54,365
June 1
11
69
73
18,000
35,200
1898
Apr. 26
57
696,000
May 17
64
195,200
June 14
82
432,800
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1894
1895
1896
July 30
July 6
July 10
" 28
82
81
80
82
1,908
231,504
90 , 000
71,000
Aug. 1
" 21
Aug. 8
79
82
86
6,350
117,513
39,200
Sept. 12
79
19,272
1897
1898
July 21
81
172,000
Aug. 24
Aug. 2
" 23
78
78
82
230,400
288,000
96,000
Sept. 14
Sept. 27
83
73
50,000
238,400
205
PULSES OF POLYARTHRA PLATYPTERA continued.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1894
1895
1896
1897
Oct. 17
Oct. 23
58
51
1,140
408
Nov. 27
33
74,942
Dec. 18
Dec. 29
Dec. 14
39
35
40
21,147
37,560
7,300
Oct. 5
71
816,000
Nov. 15
47
22,400
1898
Oct. 11
" 25
65
49
47,500
37,500
Nov. 22
40
6,000
Dec. 20
33
63,400
An examination of this table and the graphic presentation (PI.
V.) of the seasonal distribution will show at once the uniformly
small numbers attained at low temperatures. Between October
15 and April 15, that is below 60, no pulse exceeding 100,000 is
reached save one of 122,400, February 21, 1899, at 33. Of all the
records in this period only seven exceed 50,000. On the other hand,
during the warmer months, above 60, the pulses have a much
greater amplitude. Four of them exceed 400,000, and there are
twenty-two records above 100,000. The summer pulses are often
separated by minima which approach midwinter levels, but in spite
of this the general level of summer occurrences is much higher than
that of the colder season. In 1898 the average from April 15 to
October 15 was 30,861 per m. 3 , and for the other months of the year,
15,813, or about half the number in the warmer season. From
these facts of distribution it is apparent that though perennial the
species finds its optimum conditions at temperatures above 60. The
statement of Hempel ('99) that it thrives best in cold water is not
borne out by the statistical examination in any of the years.
The recurrent pulses of this species vary greatly in amplitude.
The largest pulse recorded was that of 816,000, October 5, 1897, at
71. It appeared in a period of prolonged low water and at the
close of one of high temperatures continued beyond the usual
September limit (Pt. I., PL XL), in a very unusual development of
Carteria and the smaller algae of the water-bloom (PL II.). Similar
autumnal pulses do not appear in other years, the autumnal develop-
ment as a rule not exceeding to any noticeable degree that of mid-
summer. There has been in every fully tested spring a large vernal
pulse, usually at the time of the spring volumetric maximum, or
thereabouts. In 1896 and 1898 it was the largest pulse of the year.
206
This was not true in other years, but collections in those years were
too infrequent to trace the seasonal distribution of the species with
accuracy at that season. It is volumetrically of some importance
in determining the quantitative fluctuations in the total plankton.
Computations based on its average size indicate that approximately
600,000, including eggs, would be required to form 1 cm. 3 of plank-
ton. On this basis, and allowing 10 per cent, for interstices, it
constituted at the time of its vernal maximum in 1898 about 10
per cent, of the total volume of the plankton (silk-net catch).
The table on pages 204 and 205 lists 43 pulses, of which 6 lie out-
side of the period included in Plates I. and II. Of the 38 remaining
pulses' 16 coincide in location with the whole or a part (in case of
divided culminations) of the pulses of the chlorophyll-bearing organ-
isms; 12 follow at the next collection, usually at intervals of one
week ; and 6, after a fortnight. The remaining 4 do not bear this rela-
tion, occurring in autumn or midwinter, when all pulses were feeble
and ill-defined. A comparison of Plates I. and II. with V. will
show that not all of the chlorophyll-bearing pulses are attended by
pulses of Polyarthra; nor is there any constant relation, excepting
the vernal pulse, between the size of the pulses of the two groups
of planktonts in question. Nevertheless, the dependence of the
recurrent periods of rapid multiplication of Polyarthra upon the
rhythmic occurrences of the chlorophyll-bearing organisms upon
which they largely depend for their food is strongly suggested by
the data here offered. Food relations thus dominate the repro-
ductive cycles.
The pulses of Polyarthra form a considerable portion of many of
the pulses of the total Ploima, and it is but natural that we should
find a coincidence in their locations. This may be followed for 1898
in Table I . In a number of instances the culminations of the pulses
are not exactly coincident, but separated by the interval between
two collections. The association of the two pulses is, however,
apparent in every case, and a similar relation may be traced in prior
years.
These recurrent pulses afford evidence for the polycyclic habit
of this species. Additional proof of this phenomenon is found in the
evidences of sexual reproduction either male or winter eggs
attached to the female which have attended many of the pulses.
The eggs of this species, both summer and winter forms, are very
207
readily detached in the manipulation of the plankton, so much so
that in 1898 less than 6 per cent, remained attached. More or less
uncertainty attends the determination of the parentage of detached
winter and male eggs, so that decisive proof of sexual reproduction
is best obtained from the attached eggs. In Table I. will be found
the records of free and attached male and winter eggs recorded in
1898. Evidence will be found in this of sexual reproduction at-
tending the pulses of March, April, May, September, and December.
The presence of winter eggs at intervals throughoutrthe greater part
of the year may be due either to their continual production or, as
seems more probable, to their continuance in the plankton for some
time after their formation. The presence of attached winter eggs,
or of larger numbers of free winter eggs, seems to mark the culmina-
tion and decline of the pulse. Male eggs, on the other hand, are
more generally present during both the rise and decline of the pulses.
Somewhat similar evidence of sexual cycles attends many of the
larger pulses in years prior to 1898.
This species affords a striking example of a perennial eulimnetic
planktont. It is found in midwinter under the ice in water at the
freezing point, and even under these conditions it multiplies, pro-
ducing pulses whose amplitude surpasses that of many rotifers of the
plankton, and runs a reproductive cycle similar to, though of less
amplitude than, those at other seasons of the year. It shares with
other organisms the vernal outburst, and repeats the process in
summer months under maximum conditions of heat and in waters
whose chemical condition is very different from that in which the
hiemal and vernal pulses appeared. Successive generations of this
species are thus adapted to widely different conditions. Through
all the changes incident to ice, stagnation, flood, sewage pollution,
changing temperature, the wax and wane and change of food, the
constant and unceasing warfare of enemies which prey upon it and
of parasites which plague it, and, above all and continuously, the
removal of countless individuals from the place of their origin by
the ceaseless current of the stream, this species lives on, holds its
own in the plankton, and repeats year after year the same sequence
of rhythmic pulses of occurrence in the river water. The secret of
the process doubtless lies in its capacity to produce repeatedly these
crops of winter eggs which serve to seed the environment and start
208
anew the cycle of growth and reproduction whenever the favorable
conditions prevail.
There is in this species no hard lorica whose variable processes
might serve to demonstrate to every observer its capacity for varia-
tion. This is doubtless one of the reasons why we do not find a host
of new species and varieties oiPolyarthra as in the case of Brachionus.
It is subject to considerable variation in size, and the swimming
lamellae vary in length, width, and serrations. Hempel ( '99) records
Wierzejski's var. euryptera in our plankton, and I have often
observed it, but no record was kept of it since the characters which
define it are not readily seen in plankton enumeration. Weber ( '98)
has mentioned, without designating by name, a long-spined variety
which I find very common among the individuals which occur in
the Illinois.
This planktont is subject to attacks of internal parasites (Sporo-
zoaf) which infest it at the times of its maximum pulses, though
never to the extent observed in the case of Bimcerium in Brachionus.
It is very frequently loaded down by Colacium, and some of the
smaller peritrichous Ciliata are often found upon it. The absence
of a hard lorica has served to obscure somewhat its food relations
to whatever animals prey upon it.
Polyarthra platyptera is a cosmopolite, and is apparently found
generally in the fresh-water plankton. Jennings ( '00) reports it as
abundant in the waters of the Great Lakes, and it has been found
generally in American waters. Zacharias ('98) and Marsson ('00)
find it in pond and stream waters of Germany; Stenroos ('98)
reports it as a predominant rotifer in the plankton and littoral regions
of Finland waters ; and Borge ( '00) finds it in Swedish plankton. It
has also been found to be an important constituent in the plankton
of European streams. Skorikow ('96) finds that it is the most
abundant rotifer in the summer plankton of the River Udy, consti-
tuting almost a third of the total rotifers. There are indications
in his records of recurrent pulses, and the largest numbers are found
in September. Zimmer ('99) finds it perennial in the Oder, but
never abundant. Schorler ( '00) finds it in the Elbe from April to
September, with maximum in August. Lauterborn ('98a) lists this
species among the perennial rotifers, and states that it is dicyclic in
the Rhine and its adjacent waters, which he has examined quite
thoroughly. The vernal sexual period begins with the appearance
209
of the male eggs in March, and winter eggs follow in April and May.
The second sexual period extends from the end of July to the end
of October, with a maximum in September-October. This bears
some resemblance to the distribution in the Illinois, with the
exception that the recurrent cycles which make the species poly-
cyclic were not noted, and that male or winter eggs were not present
in the colder months. It may be that the application of the
quantitative statistical method with brief intervals of collection in
the Rhine would reveal a still closer correspondence in the seasonal
routine of Polyarthra in the two streams. Wesenburg-Lund ('98)
finds that temperature has nothing to do with the appearance of
the sexual cycle of this species in Danish waters. Males were
found in December, as also (eggs only) in the Illinois. He also
found differences in different bodies of water as to the times of the
sexual cycles. Apstein ('96) has found this species perennial and
one of the most abundant rotifers in plankton of the lakes near
Plon, Germany, with maximum period from April to August, and
in November in one lake, and in July-August in another. The
sexual cycle was noted in May- June only. Seligo ('00) finds the
species perennial in lakes near Danzig, with large numbers in April
and July. His collections were too widely separated to trace fully
the seasonal fluctuations. Burckhardt ('OOa) finds Polyarthra in
small numbers in winter months in the plankton of Swiss lakes, and
in larger numbers in the summer, but does not trace their seasonal
fluctuations.
Pterodina patina Ehrbg. Average number of females, 37. With
two exceptions all the records of this species lie between the last of
May and the first of October. There are but four records below
70. This indicates optimum conditions for the species during the
period of maximum heat, and further evidence of this lies in the
occurrence of the larger numbers during this period. Appearances
in January-March suggest a perennial habit ; and small and irregular
numbers, that the species is largely adventitions. Hempel ('99)
also records P. valvata Hudson from Quiver Lake.
Rattulus tigris O. F. Mull. Average number of females, 207. I
have not found this species in any year later than October, though,
as shown in Table I. , it appears in January at minimum temperatures,
and continues in small numbers and somewhat irregularly until
autumn. These conditions and the absence of pulses suggest that
210
the species is adventitious in the plankton. The greater part of the
occurrences were recorded above 50 and the larger numbers above
60, indicating an optimum during summer months. The record in
Table I. refers to the species figured by Jennings ('00) under this
name.
Rattulus sulcatus Jennings was found seven times in the plankton
in July and August during maximum temperatures. It is probably
adventitious in the plankton.
Salpina brevispina Ehrbg. was found September 5, 1895, at
74, and April 29, 1896, at 70.
Salpina eustala Gosse was found July 13, 1894, at 82.
Salpina macracantha Gosse was found September 5, 1895, at 74.
Salpina ventralis Ehrbg. was found July 29, 1895, at 75. In
common with other species of the genus it is adventitious in the
plankton.
Schizocerca diversicornis v. Daday. Average number of females,
46. The earliest record of this species was June 1, 1896, at 70 ; and
the latest, September 20, 1895, at 78. Most of the records and
the larger numbers are in July-September during the period of
maximum heat, in which its optimum conditions must be found.
Egg-bearing females were also found in these months. This species
is closely related to the Anuroza aculeata group, and like it is exceed-
ingly variable, especially in degree of development of the various
spines. Variety homoceros Wierz. was found in May, June, and
August, 1896. Five sixths of all the individuals recorded were
found in 1896, and the fact that this was a year of unusually dis-
turbed hydrograph (Pt. I., PI. X.) suggests that this form may be
to some extent adventitious in our plankton, but no direct relation
to the access of flood waters can be traced.
Lauterborn ('98a) lists this species among the summer planktonts
of the Rhine, and Seligo ( '00) finds it in large numbers, with a maxi-
mum in July, in lakes near Danzig. Zacharias ('98) reports it in
German pond plankton, Zimmer ('99) finds it in the Oder, and
Schorler ( '00) in the summer plankton of the Elbe.
Synchazta pectinata Ehrbg. Average number of individuals,
3,950; of eggs, 13,823. It was much more abundant in previous
years, averaging in 1897 23,227 and 28,230; in 1896, 7,064 and
7,927; in 1895, 13,071 and 4,730; in 1894, 7,520 and 1,659. The
effect of the disturbed hydrograph of 1896 is seen in the smaller
211
numbers of that year, while the larger numbers in 1897 may be
attributed to the more stable conditions. The small numbers in
1898 do not seem to be correlated with any feature of the environ-
ment.
This species has been found in every month of the year, and is
thus perennial in our plankton. As will be seen, however, in Table I. ,
the most of the occurrences and a much greater proportion of the
individuals are found between May and October, and thus above
60. The same limitations are found in the other years, with the
exception that in 1896 there was a more continuous and larger de-
velopment from the last of February. In the table which follows
it may be noted that all of the pulses but four are at temperatures
above 70, and of these four none exceeds 25,000, and two do not
exceed 2,500. The optimum conditions for the species in our
waters are therefore above 70. The average temperature at the
time of the larger pulses is near 80. The vernal pulses are poorly
defined, as are likewise the autumnal ones. It is a midsummer
species in our waters, with its maximum in August.
PULSES OF SYNCH^ETA PECTINATA.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1895
1896
Mar 3
35
6 360
1897
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1894
Tiilv 1 3
83
74 606
1895
1896
July 23
Tnlv in
80
80
1,749
22 200
Aug. 12
85
7ro
175,230
Sept. 12
79
27,740
" 28
82
38,000
1898
1898
July 19
84
350
20,800
" 24
Aug. 2
" 23
78
78
82
264,000
12,000
3,200
Sept. 27
73
30,400
(IS)
212
Of the 18 pulses listed in the preceding table 17, fall within the
limits of periods included in Plates I. and II. Of these 17 there are
7 which coincide with, and 9 which follow shortly after, -the culmina-
tion of the pulses of the chlorophyll-bearing organisms, while 1, a
small one in March, 1896, shows no such correlation. Food is thus
a primary factor in the production of these recurrent pulses. -As
will be seen in Table I., these pulses uniformly coincide with those
of the total Ploima, and a similar relation may be followed in pri< >r
years.
The eggs of this species are not usually carried by the female for
any length of time, and are rarely found attached in preserved
material. For this reason the sexual cycles are not easily followed
with accuracy in the statistical data. It may be seen in Table I. that
the free winter eggs belonging to both species of SynchcBta are most
numerous in the period of the larger pulses, and that their occur-
rences show some tendency to coincide with these pulses. Proof
that these pulses terminate in sexual reproduction is thus lacking,
though it seems probable from some of the evidence.
Synch&ta pectinata has not been widely reported from American
waters. Jennings ('94) finds it in Michigan and Kellicott ('97) in
Lake Erie, but it has not been elsewhere reported in American
plankton. It appears, however, in many European records. Skori-
kow ('96) finds it in the summer plankton of the River Udy, in
Russia ; Zimmer ( '99) finds it in common with 5. tremula in the Oder
throughout the year. He makes the statements that it is never
rare, is somewhat more abundant in the spring, and is, at other
times, present "in relativ gleichmassiger Haufigkeit." In the
light of our results it seems probable that the data at Zimmer 's
disposal were insufficient to justify his conclusions as to the uniform-
ity of its seasonal distribution. Schorler ('00) finds it in the Elbe
in April, May, and October, with a maximum in May. Lauterborn
( '98a) finds it perennial in the plankton of the Rhine, and lists it
among the dicyclic species with two periods of sexual reproduction,
one in April and one from the end of July to October. Judging from
the character of the statistical data which have been presented for
this and other species in the Illinois it seems probable that the later
period noted by Lauterborn may include several cycles, and that
the species is usually a polycyclic one. Seligo ('00) reports it
perennial in waters near Danzig, with largest numbers in April and
213
September. Apstein ('96) finds that this species (including 5.
tremula and 5. grandis] is one of the most abundant in lakes near
Plon, with variable maxima in different bodies of water. He finds
it perennial in one case, and reports vernal maxima. Winter eggs
were found in March and April.
Synch&ta stylata Wierz. Average number of individuals, 120,391 ;
of eggs, 17,797. In 1897, 42,577 and 9,127; in 1896, 24,099 and
5,125; in 1895, 155,880 and 2,418; in 1894, 8,582 and 132. This
species affords an exception to the general rule hitherto observed
among the rotifers of our plankton in that it is more abundant in
1898 than in the previous year. As will be seen in the following
table both the vernal and autumnal pulses are unusually large in
1898, while in the previous year the vernal pulse is only moderate
and the autumnal pulse is scarcely to be detected. For some
reason the prolonged low water and sewage contamination of the
autumn of 1897 was not favorable to the usual grow r th of this
species. It may be that it was crowded out by the unusual develop-
ment of Polyarthra at that season (PL V.).
PULSES OF SYNCH^ETA STYLATA.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
" 25
33
3,648
330
1899
Jan. 14
34
12,000
Feb. 14
32
19,200
" 22
Mar. 21
51
37
58,000
5,600
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1896
Apr. 29
70
380,586
May 25
75
10,800
June 17
76
79,200
60
1 139 000
June 21
77
795 200
" 31
70
61,600
214
PULSES OF SYNCH/ETA STYLATA continued.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1894
1895
1896
1897
1898
Aug. 1
Aug. 8
79
86
10,287
8,400
Sept. 27
73
12,225
July 21
July 19
81
84
103,200
64,800
Sept. 7
Sept. 27
80
73
28,000
265,600
Aug. 2
" 23
79
82
170,400
24,800
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1 894
1895
1896
1897
Oct. 17
58
63,935
Nov. 27
Nov. 17
Nov. 9
" 30
33
44
50
34.5
901,901
114,000
26,400
87,200
Dec. 11
32
1,121,056
Oct. 5
" 19
71
65
12,000
15,800
Dec. 14
36
72,200
1898
Oct. 25
49
824,500
Nov. 15
41
110,000
Dec. 6
20
34
33
42,500
59,200
This is the most abundant of all the rotifers in our plankton,
exceeding by 30 per cent. Polyarthra, the next in abundance. It
constituted one fifth of the total Ploima in 1898, and is accordingly
a large factor quantitatively and ecologically in the economy of the
plankton of the Illinois River.
It is a perennial planktont, occurring in six sevenths of our
collections and usually in considerable numbers. The distribution
in 1898 (Table I.) is a fair index of the usual seasonal routine, with
the exception that in all prior years the July-August minimum is
more pronounced and better sustained. The development in
January-February is never large, rarely exceeding 20,000. In
March, numbers rise rapidly, usually with a minor pulse, the re-
covery from which in April culminates in a vernal pulse, which in
three of the six years was the largest of the year. Following this
vernal pulse there is a series of smaller pulses throughout the sum-
mer. The decline of the June flood, when this occurs, seems to offer
favorable conditions (cf. foregoing table and Pt. I., PI. IX.-XII.)
for the development of a pulse which is but little smaller than the
vernal one. It may be of some significance that this pulse and the
215
vernal one both occur on the decline of the major floods of the
year, and that the relative proportions of the two floods are to some
degree paralleled by the amplitude of the pulses of Synch&'ta which
attend their decline. The effect of the impounding backwaters as
reservoirs for the greater development of the plankton is suggested
by these data.
Following the midsummer minimum is an autumnal pulse whose
amplitude and location alike are subj ect to much variation. As will be
seen in the table on pages 213 and 214, the maximum jmtumnal pulse
is located twice in October, twice in November, and once in Decem-
ber. This may be due to the fact that the collections are insufficient
in some of the years, or to the probability that any one of several
recurrent autumnal pulses may be the major pulse of that season.
An examination of the seasonal distribution. in 1898 (Table I.)
and of the location and temperatures of the pulses recorded in the
table on pages 213 and 214 will suffice to demonstrate the capacity of
this species to develop at all temperatures within the seasonal range.
The largest pulse (1,139,000 on May 3, 1898) is at 60, and the next
in size (1,121,056 on December 11, 1895) is at 32. It will, however,
be seen in the two tables that the pulses and the numbers in general
during the periods of maximum heat and cold are not so large as in
the intervals of more moderate temperatures. The impetus of the
autumnal development may carry some of the pulses over in to
minimum temperatures, but the level of development declines
thereafter. There is thus something of a tendency for the average
temperature of the larger occurrences to approach the average
temperature of the year.
The number of pulses listed in the table on pages 213and214is38.
Of these, 34 fall within the period included in Plates I. and 1 1. of the
pulses of chlorophyll-bearing organisms. Of the 34 there are 18
which coincide in location with these plant pulses, 12 which follow
at a brief interval, and 4 which bear no such relation, three of the
last being minor winter pulses.
The dependence of the recurrent periods of rapid multiplication
of Synchata the most abundant rotifer of the plankton upon the
rhythmic increase of the food supply is thus fairly demonstrated.
The coincidence of the pulses of Synch&ta with those of the total
Ploima is readily seen in Table I., and is equally apparent in prior
vears.
216
Eggs of this species are not carried by the parent for any length
of time, so that reproductive cycles are not easily traced. The total
number of the summer eggs of Synchceta will be found (Table I.) to
fluctuate somewhat with the pulses of the species. The free winter
eggs, belonging probably to both species of Synchatta, also show
some tendency to predominate at and after the culmination (Table
I.) of the pulses. A female carrying a male egg w T as recorded during
the rise of the spring pulse in 1898, and attached winter eggs were
noted at the vernal pulse in 1895 and 1897. The evidence points
toward the culmination of these pulses in a sexual cycle.
The soft and flexible nature of this rotifer and the absence of
spinous outgrowths have made whatever variability the species
possesses less evident than it is in such a genus as Brachionus. There
is considerable variation in size possibly due to age even in the
same collection. The determination of preserved material of this
genus is fraught with insuperable difficulty. The separation of
pectinata and stylata in our records is at the best only probable. It
may be that other species of Synchceta have been included w r ith the
individuals referred to stylata. In any event the result of the
division has led to symmetrical results comparable with those of
other planktonts. Synch&ta is often parasitized at the times of the
larger pulses by some sporozoan (?). At the maximum of the
vernal pulse in 1898 over 4 per cent, of the individuals were thus
affected, the infestation continuing through the decline of the pulse.
External parasites, Colacium and Rhabdostyla, are rare.
This species has not been found widely in the plankton, possibly
because of the confusion of stylata, tremula, and pectinata in identifi-
cation. From the large numbers reported in almost every instance
where it has been found, the expectation of its wide-spread occur-
rence is at least raised, waiving in this connection the possibility of
specific confusion. Jennings ( '94) found it to be very abundant in
towings in Lake St. Clair, and ( '96) in Lake Michigan near Charle-
voix. He finds it less abundant in the summer plankton of Lake
Erie ('00). Stenroos ('98) reports it as one of the most abundant
limnetic rotifers in Lake Nurmijarvi in Finland in the summer, and
Skorikow ( '97) finds that next to Polyarthra it is the most abundant
rotifer in summer months in the River Udy near Charkow, Russia.
His figures of occurrence show some traces of recurrent cycles in
these months, with maximum numbers at the first of August. Lau-
217
terborn ('98a) lists it among the summer rotifers of the plankton
of the Rhine. The genus is in need of a thorough revision in the
light of possible variation.*
Taphrocampa annulosa Gosse. Average number, 71. Found
in September, at 73. Evidently adventitious.
Triarthra longiseta Ehrbg. Average number of individuals,
3,147; of eggs, 293. This species was about twice as abundant in
the stable conditions of 1897, and was present in less than half these
numbers in the recurrent floods of 1896.
It is a perennial species, having occurred in every month of the
year. The continuous occurrences and the larger numbers lie in
all years between May and October and above 60. In 1898, only
about 3 per cent, of the total individuals were found below this
temperature. With the exception of the vernal pulse of 1898 all
of the larger numbers were found in the period of maximum heat.
The optimum conditions for this species are thus found within that
period and above 70.
The seasonal routine of the species is varied somewhat from year
to year. There is usually a slight vernal pulse larger than usual
in 1898 and this is followed by recurrent pulses throughout the
summer. The season closes without a predominant autumnal pulse,
and after September the numbers fall and the occurrences become
sporadic until the following April.
The pulses of this species are listed in the following table, which
gives their locations and temperatures.
Of the 21 pulses recorded, 18 are within the periods of the plant
pulses shown in Plates I. and II. Of these 18 there are 8 which
coincide with these plant pulses, 9 which follow after a short interval,
and 1 which shows no such relation. The dependence of the pulses
of Triarthra upon food conditions is suggested. The pulses of
Triarthra will be found on examination of Table I. to coincide in
1898 in the main with those of the total Plointa.
The pulses are never very large, and the evidences of reproduc-
tion are not well defined. Attached summer eggs attend the larger
pulses, and free winter eggs of the species were found in October-
November in 1898. In previous years free or attached eggs attended
vernal or summer pulses at times. The evidence indicates a poly-
cyclic habit.
* See Rousselet, '02.
218
PULSES OF TRIARTHRA LONGISETA.
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
7n
73
4 000
" 27
80
6,000
31
70
1,000
Year
Date
Temp.
No.
Date
Temp.
No.
Date
Temp.
No.
1895
July 18
80
19,080
Aug. 21
82
10,683
Sept. 12
79
2,336
1897
July 21
81
49 , 600
Aug. 17
79
9,600
Sept. 7
80
70,000
1898
July 26
89
28,000
Aug. 30
83
6,400
Sept. 27
73
14,400
This is an exceedingly variable species. It varies in the relative
length of the three long setae, in their spinosity, and in the location
of the posterior one. Many of the individuals in our waters resemble
the form described by Plate ( '85) as T. terminates. The long-spined
form described by Zacharias ( '94) as var. limnetica is also abundant.
It is doubtful if either form is worthy even of varietal distinction.
This species has been reported only from Lake Erie and the
Illinois River in this country, and seems to be rare in the former.
Weber ('98) finds it abundant in the plankton of Lake Leman;
Burckhardt ('00 and 'OOa) reports it as wide-spread and almost
perennial in Swiss lakes, but with its maximum in December-
February, and slight development during warmer months. Borge
( '00) finds it to be one of the common rotifers in the summer plank-
ton in Sweden ; Marsson ( '00) reports its perennial seasonal range in
several German waters, with greater numbers during the warmer
season. Apstein ('96) gives it a perennial distribution in Lake
Plon, with larger numbers in June-November, and maximum in
June- July or August. According to Seligo ( '00) the species is per-
219
ennial in lakes near Danzig, rivaling Polyarthra in abundance, and
exhibiting maxima in the warmer months from April to October.
It is also a member of the potamoplankton of European streams.
Skorikow ('97) finds it in summer months in the Udy, and Zimmer
('99) reports if as present in small numbers and irregularly in the
Oder from April to November. Schorler ('00) finds it in the Elbe
in May-October with maxima in May and September, and Lauter-
born ( '98a) includes it in his list of perennial rotifers in the plankton
of the Rhine. It has two sexual periods, the first in March-May
and the second in July-October, and he suggests the probability of
a poly cyclic habit in some waters.
Trochosphczra solstitialis Thorpe was found June 27, July 2, and
August 15, in 1896 ; in 1897, on May 25 and July 14-30. Free winter
eggs were taken August 15, 1896. All occurrences were above 66.
These records were all from plankton taken in mid-channel of the
main stream. Trochosph&ra was found in greatest abundance at
the outlet of Flag Lake (Pt. I., PI. II.) in July, reaching 9,664 per
m. 3 at 72. It was also found in August in the weedy backwaters
of Dogfish Lake. Both of these backwaters connect with the river
(Pt. I., PI. II.) below the point at which our collections were made.
It was either introduced from some similar backwater higher up the
stream than our plankton station, or developed in the river itself.
SCIRTOPODA.
This order is represented in the plankton by a single species,
whose discussion will suffice for the order.
Pedalion mirum Huds. Average number, 4,524. This is a
summer planktont of somewhat definite temperature limits. The
following table combined with the data in Table I. will suffice to
characterize its seasonal fluctuations.
Its limitation to temperatures above 60, indeed almost 70, is
apparent. There are in all but two records below 60, and but four
below 70. It is a typical midsummer planktont, with several
recurrent pulses during the period of maximum temperatures.
The location of these pulses with reference to those of the
chlorophyll-bearing organisms is significant. As shown in Table I.,
they follow immediately, or coincide with, those of the synthetic
organisms. For example, the apices of the pulses of Mastigophora,
220
Year
First record
First maximum
Date
Temp.
Date
Temp.
No.
1894
June 29
July 6
July 28
July 21
July 26
83
80
80
84
89
2,592
330,932
20,000
80 , 000
99,600
1895
1896
May 25
June 28
June 21
70
75
77
1897
1898
Year
Second maximum
Last record
Date
Temp.
No.
Date
Temp.
1894
Sept. 17
Oct. 2
Sept. 16
Sept. 14
Nov. 1
72
63
71
73
45
1895
Aug. 21
Aug. 15
Aug. 17
Aug. 16
81
81
79
77
3,561
77,600
79,200
22,400
1896
1897
1898
Bactilariacea, and Chlorophycecs in the period in question in 1898
are (PI. II.) July 19, August 9, August 30, and September 27. The
apices of the Pedalion pulses are July 26, August 16, and September
27, the last coinciding with the pulse of chlorophyll-bearing organ-
isms. In 1897, the intercalation of the two pulses is apparent, and
in 1896, two out of three pulses are intercalated and a third is
coincident. As will be seen in Table I., these pulses of 1898 are
approximately coincident in many cases with those of other roti-
fers Synchceta, Polyarthra, Triarthra, and Brachionus. The sig-
nificance of this intercalation lies probably in the food relations of
the two groups of organisms.
Females with a single egg attached to the body have been noted
at the times of the maxima of the pulses, or immediately thereafter,
221
in five instances. On the pulse of July 26, 1898, a female with four
male eggs was found.
This species was not reported by Apstein ('96) from the lakes
of Holstein, but was found by Lauterborn ( '98a) in the Rhine and
its backwaters. Here also it was a summer form, appearing about
the middle of June, with a maximum in August or September and
disappearing late in October, conditions of distribution much re-
sembling those in the Illinois. It is regarded, along with other
summer forms, as monocyclic. The appearance in- our waters of
male eggs July 26, at the height of the first pulse, leads to the in-
ference that there may be several cycles; for example, three in
1898, with the recurrent pulses, in a single summer season. Weber
( '98) gives it as a summer rotifer in Switzerland, and Skorikow ( '97)
finds it in July-September in the Udy River, in Russia ; but it is not
reported from the Oder by Zimmer ('99), nor from the Elbe by
Schorler ('00). Kellicott ('97) finds it in Lake Erie in small
numbers in the summer.
In addition to the species of rotifers noticed above, Hempel ('99)
has reported the following in the Illinois River or its backwaters:
Flosculana ornata Ehrbg., Limnias ceratophylli Schrank, Cephalosi-
phon limnias Ehrbg., CEcistes intermedius Davis, 0. mucicola Kell.,
Pedetes saltator Gosse, Furcularia forficula Ehrbg., F. longiseta
Ehrbg., Eosphora aurita Ehrbg., Diglena grandis Ehrbg., D. catellina
Ehrbg., D. biraphis Gosse, Ccelopus tenuior Gosse, Scaridium longi-
caudum Ehrbg., Distyla gissensis Eckstein, D. okioensis Herrick,
D. stokesi Pell, and D. hornemanni Ehrbg.
GASTROTRICHA.
Ch&tonotus sp. occurred singly in the plankton August 29, 1896,
July 30, 1897, and February 15, 1898, with a temperature range of
32.5 to 84.
ENTOMOSTRACA.
Average number, 47,042. In 1897, a more stable year, 91,050;
in 1896, a year of disturbed hydrograph, 50,158; in 1895, in more
stable conditions, 148,348. The Entomostraca appear in every collec-
tion at all seasons of the year. The decline to the winter mini-
222
mum occurs in November-December. Numbers are at a minimum
(generally less than 5, 000 per m. 3 ) in midwinter (January-February) ;
rise in March to about 25,000 per m. 3 ; and attain the maximum for
the year in a vernal pulse of 200,000 to 1,500,000 in April-May.
Following this, there is frequently a second pulse of large proportions
in June, which in 1898 exceeds (Table I.) that of May. During the
remainder of the year there is usually a series of recurrent pulses, of
declining amplitude in 1896 and 1898, but rising to unusual heights
(618,750 on September 9) in the stable conditions of 1897. In
the main the pulses of Entomostraca coincide with or approximate
to the location of those of the other organisms of the plankton, and
often show correlations in amplitude.
BRANCHIOPODA.
Eubranchipus serratus Forbes. Young branchiopod larvae
questionably referred to this species appeared in the plankton
in January-March, 1899, in small numbers at minimum tempera-
tures.
CLADOCERA.
Average number, 6,068 per m. 3 In 1897 they were more abun-
dant, averaging 17,863 per m. 3 in the more stable conditions of that
year. In 1896, a year of recurrent floods, numbers fell to 7,7 19, while
in 1895, a year of low water in spring, when many of the Cladocera
attain their maximum, the greatest average, 31,937, was recorded.
The phenomenal number of 443,716 per m. 3 appeared on June 19 in
the stable low water (1.80 ft.) then prevailing. In 1894, another
year of low levels, the annual average was also large (23,952), though
probably enhanced by the fact that collections were not made in
flood waters in this year.
The Cladocera appear in all but 10 of the 182 collections enu-
merated, the ten exceptions falling in November (1), January (2),
February (6), and April (1), and usually in flood waters or, as
in 1895, in stagnation conditions under the ice. Although the
Cladocera occur in all months of the year, they nevertheless, as a
group, exhibit decided temperature adaptations, as appears from
the fact that all records in excess of 4,000 per m. 3 fall between May 1
and September 1 with but 6 exceptions, 4 in the phenomenally
223
early spring of 1896, and 2 in the delayed high temperature of
October, 1897.
The minimum records (less than 500 per m. 3 ) are found during
minimum temperatures. The numbers increase slightly (generally
less than 2,000) as temperatures rise in March- April, rise abruptly,
as they approach or pass 70, to a vernal maximum in May- June,
and decline during midsummer excepting when unusual pulses of
Moina or Diaphanosoma raise the level of the pulse maxima above
25,000. This decline continues in channel plankton through the
autumn until the low level of approximately 2,000 per m. 3 , at the
most, is again attained in October, and falls irregularly to 500, or
less, as minimum winter temperatures arrive in December. Ex-
ceptions appear in 1897, when a well-defined autumnal pulse of
large amplitude (193,500) is found on September 14, and is followed
by others of declining amplitudes (137,600, October 5; 5,520, No-
vember 15 ; 4,240, December 14) during stable autumnal conditions.
All of the records above 4,000 per m. 3 , with one exception, are
found at temperatures above 45, and all in excess of 8,000, with 4
exceptions, after the vernal rise in temperature passes 70 in April-
May, and before the autumnal decline reaches this point in Septem-
ber. The Cladocera are thus planktonts of the warmer channel-
waters.
The relation which hydrographic conditions bear to the seasonal
occurrences of Cladocera is apparent in the yearly averages above
quoted, and appears still more clearly in a comparison of the
cladoceran population and movement in river levels in July-
December, 1897 and 1898, as given below.
Average No.
Cladocera
per m. 3
July
August
Sept.
Oct.
Nov.
Dec.
1897
1898
1897
1898
1897
1898
1897
1898
1897
1898
1897
1898
12720
3050
13960
3756
70675
1700
40350
1615
2532
620
1945
236
Total movement
in river levels,
in ft.
5.2
7.4
2.6
7.5
0.6
6.2
0.6
3.9
2.2
2.6
0.5
2.4
224
Hydrographic changes affect the Cladocera by increasing the
amount of silt and flocculent debris in suspension, which, by ad-
herence to the swimming antennas and flotation processes of the
animal, tend to impede its movements and sink it to the bottom,
where it is removed from its normal feeding area and readily becomes
the prey of the larger organisms of the bottom fauna. Barren flood
waters also tend to displace and wash away in the increased current
the Cladocera which have developed in the stream, and to afford
both less food and less time for their further development.
The occurrences of the total Cladocera fall into the type of
recurrent pulses, though with slightly less distinctness than in the
case of individual species of the group. Such pulses can be traced
in all seasons in which records were made at short intervals, and
suggestions of their occurrence appear in the less frequent records
of other seasons. Thus in July-December, 1897, (PI. IV.), there
are 6 well-defined pulses culminating at intervals of 3(1), 4(2), 5(1),
and 6(1) weeks. In 1898 (Table I.) the pulses are less regular in
the flood waters of the disturbed year. In 1896, when records were
frequent, we can trace pulses in March, May, June, July, August, and
September. The character of these pulses is well illustrated in the
vernal pulse of 1898 (Table I. and PI. IV.), culminating June 7 at
136,000. The species which share in this pulse are Alona affinis,
A. costata, A. quadrangularis , Bosmina longirostris*, Ceriodaphnia
scitula*, Chydorus sphcericus* , Daphnia hyalina*, D. cucullata*,
Diaphanosoma brachyurum, Leptodora hyalina, Macrothrix laticornis,
Moina micrura, Pleuroxus denticulate s, Scapholeberis mucronata, and
Simocephalus serrulatus. Of these, only the five marked by the
asterisk occur in numbers sufficient by our methods to delineate a
pulse. The other species are accordingly of little consequence in
modifying the form or location of the pulse. The June volumetric
pulse (Part I., PI. XII.) culminates June 14 at 6.99 cm. 3 per m. 3 ,
though the record for June 7 is also high (5.28). The cladoceran
pulse culminates June 7 at 136,000. On this same day four of the
dominant species also reach their culmination, viz. : Bosmina
longirostris (62,800), Ceriodaphnia scitula (55,800), Daphnia cucul-
lata (3,400), and D. hyalina (11,600), the remaining 2,400 being
contributed by other species. Chydorus spharicus, which appears
this spring only in small numbers, attains its maximum (7,880) on
May 24, two weeks earlier, though the record for May 31 is also high
225
(5,040), indicating a probable maximum between these dates. In
other seasons, for example in 1896 and 1897, the maxima of this
species coincide generally with those of other Cladocera, so that this
divergence seems to be anomalous. An inspection of the table of
records for 1898 gives a remarkably uniform and coincident rise and
decline of the pulses of the several species which constitute this
characteristic vernal pulse.
No effort has been made by me to determine the total cladoceran
fauna of the Illinois River. Only those species are here given which
have appeared in our plankton enumeration. A number of others
are known to occur in the littoral fauna, and a few scattering indi-
viduals found in the plankton were not identified.
Of the 25 forms here listed, only 10 named in the sequence of
their relative numbers as shown in grand totals may be regarded
as typical planktonts, autolimnetic in channel plankton-, viz. : Moina
micrura, Bosmina longirostris, Daphnia cucullata and vars. upicata
and kahlbergiensis, D. hyalina, Ceriodaphnia scitula, Chydorus
sphcericus, Diaphanosoma brachyurum, and Leptodora hyalina. Of
the ten, the last named and the varieties of D. cucullata appear to
be of little quantitative importance in the channel plankton, though
it may be that our methods of collection fail adequately to represent
Leptodora. Of the remaining 15 species, Alona a~ffinis, Ceriodaphnia
reticulata and C. rotunda, Scapholeberis mucronata, and the two
species of Simocephalus are the only adventitious Cladocera of
quantitative importance, and this only to a relatively small extent.
DISCUSSION OF SPECIES OF CLADOCERA.
Alona affinis Leydig. Average number, 36. This species has
a well-defined seasonal distribution. It appears in autumn in the
last of October, as temperatures approach 40, and remains until
the end of June, when the summer maximum of 80 is re-established.
The numbers are too small (Table I.) and irregular to define its
seasonal fluctuations, though there are suggestions in the records
of late autumnal and of vernal pulses. Egg-bearing females were
recorded in January-February at minimum temperatures. No close
dependence on hydrographic fluctuations is apparent to account for
their occurrence in the plankton.
Alona costata Sars. Average number, 11. Only a few scattered
occurrences of small numbers. Earliest autumnal record, Novem-
ber 22, at 40; latest vernal, May 24, at 73.
226
Alona quadrangularis O. F. Miill. Average number, 5. A few
scattered occurrences in March-May.
Alona spp. It is probable that some of the foregoing species of
Alona are here included. There are 16 occurrences, scattered
through all months but January, April, and November, with no
large numbers and no marked seasonal distribution.
Bosmina longirostris O. F. Mull. Average number, 2,441, of
which 1,527 are adult females without large embryos, 390 with
them, and 524 immature.
I include in this species B. cornuta Jurine, for I am unable to find
any constant line of demarcation between these forms. The
longirostris form is the dominant one in the channel plankton, the
cornuta form being relatively rare.
Bosmina is a perennial planktont in our channel plankton, but
occurs in small numbers only in October-May, no record in this
period with the exception of that of October 5, 1897 (20,400), at
71, exceeding 5,000 per m. 3 , and most of them falling below r 2,000.
The records in November-March, with the exception of November-
December, 1897, all fall below 1,000 per m. 3 In like manner the
percentage of collections containing Bosmina in December-April is
lower than that in the summer, the percentages being 64, 16, 26, 47,.
and 55 per cent, respectively for these colder months, and averaging
82 per cent, for the rest of the year. The percentage of occurrences
in October-November remains high (82 and 81 per cent.), though
the numbers per m. 3 fall off greatly.
The usual seasonal distribution is as follows : In January-March
the occurrences are scattered and irregular and the numbers very
small less than 500 per m. 3 Toward the close of April the vernal
increase makes its appearance, continues slowly through May, rarely
attaining more than 5,000 per m. 3 , and at the end of this month or
early in June reaches the maximum development of the year in a
vernal pulse of 40,320 (1896) or 62,800 (1898) per m. 3 From this
summit there is an abrupt descent in a period of exhaustion to a
level of less than 2,000 per m. 3 in the last fortnight of June. During
the remainder of the year there appears a series of recurrent pulses
of less magnitude, exceeding 10,000 per m. 3 in but three instances.
These follow at intervals of four to six weeks. In July-September
the amplitude of these pulses exceeds in all cases 5,000 per m. 3 In
October (with the exception of 1897, when temperatures were un-
227
usually high), they decline in amplitude, and in November-Decem-
ber often fail to appear in the small numbers recorded. In 1894,
records are too scanty to be of significance. In 1895 there are
three well-defined pulses, and traces of a fourth in August-Novem-
ber. In 1896 there are five in May-September. In 1897 there are
six in July-December, data during the remainder of the year being
insufficient to define the pulses. In 1898 the vernal pulse in June
and a feeble one in October are the only ones which appear. The
pulses of Bosmina are best defined in the stable low water of the
last six months of 1897. During that period they closely approxi-
mate in location of maxima and minima the quantitative pulses
and those of the chlorophyll-bearing organisms and of the rotifers.
(Compare on this point the plates for 1897 in Part I. Kofoid, '03
and PI. III. and IV.) . The slopes of the pulses indicate that Bosmina
is capable of very rapid multiplication; and their coincidence with
other pulses just noted, taken in conjunction with the fact that
males and ephippial eggs appear but rarely, suggests that these
pulses of Bosmina are immediately dependent, in large part, upon
fluctuations in the food supply for their origin and for the varying
courses which they run.
The relations of Bosmina to temperature appear in the facts
that all pulses exceeding 5,000 per m. 3 in amplitude occur at tem-
peratures above 70, that the vernal rise does not proceed with any
rapidity until this temperature is attained, and that the depressing
effect of the autumnal decline below 70 is at once apparent in the
reduced numbers per m. 3 No constant relation between the pulses
of Bosmina and the midsummer heat pulses such as appears in
the records of Diaphanosoma can be traced in the occurrences of
Bosmina.
An inspection of the accompanying table, in which the mean
monthly Bosmina population per m. 3 of channel water in July-De-
cember, 1897 and 1898, is given, and also the total + and - move-
ment in river levels for these months in each year, will suggest an
intimate connection between stability of hydrographic conditions
and the increase of Bosmina. In 1897 the total movement for these
months is from five sevenths to one tenth of that in 1898, and in
every instance the Bosmina population is also greater by from 7.5
to nearly 400-fold in 1897, the more stable year. The means of the
six months are 2.03 ft. and a population of 3,691 in 1897 to 5.3 ft. and
228
BOSMINA AND HYDROGRAPHIC FLUCTUATIONS.*
Year
July
August
September
Total
movement,
in feet
Bosmina
per m. 3
Total .
movement,
in feet
Bosmina
per m. 3
Total
movement,
in feet
Bosmina
per m.^
1897
f -3.9
5
I +1.1
6,213
r -2.6
2.6\
( + o
3,973
r - -2
.6
I + -4
3,022
1898
r -6.9
7 U,
140
r -3.3
7.7
I +4.4
10
r-2.6
6
[ +3.4
IS
Year
October
November
December
Total
movement,
in feet
Bosmina
per m. 3
Total
movement,
in feet
Bosmina
per m. 3
Total
movement,
in feet
Bosmina
per m 3
1897
,(-'
I + .5
5,875
2.2\
[ +1.5
1,680
f - .6
1.2
I + .6
1,585
1898
r -1.1
3.9
1 +2.8
780
f - .6
3.2
I +2.6
32
f -2.8
3.8]
[ +1.0
60
* + = rising levels; = falling levels.
173 Bosmina in 1898. It is also true that months in which the
disparity in stability is greatest are those in which the Bosmina
ratios are greatest, and vice versa. It seems very probable that
the increased current, the lessened time for breeding, and the greater
burden of silt in flood conditions, especially rising waters, do not
conduce to the rapid increase of Bosmina in channel plankton.
The effect of the high temperatures of the late autumn of 1897
is apparent in the amplitude of the October, November, and De-
cember pulses (20,400, 3,440, and 3,440, respectively), which exceed
those of all other years at this season. Temperature thus plays
perhaps by virtue of its relation to the food supply an important
229
part in the seasonal delimitation of the amplitude of Bosmina pulses.
The Bosmina population in the plankton consists largely of
parthenogenetic females. Males and females with ephippial eggs,
were recorded only in October-December, 1897, and then only in
small numbers and isolated occurrences. Females with eggs or
embryos and the free young were found at all seasons of the year and
at all temperatures, but most abundantly at the time of the pulses.
Parasitized or fungused individuals are also found occasionally at
these seasons of greatest numbers, and the high mortality following
a pulse is evidenced by the large number of dead occurring in the
plankton. The proportions of females, females with eggs or em-
bryos, young, and* dead during the May- June pulse of 1898, may
be traced in the following records.
BOSMINA PER M. 3 , MAY-JUNE, 1898.
Date
Females
Females
with eggs
Young
Total living
Dead
Apr. 26
800
800
May 3
1,600
400
800
2,800
" 10
1,600
1,000
1,000
3,600
400
" 17
1,300
1,100
1,100
3,500
100
" 24
3,280
1,400
1,240
5,920
920
" 31
25,120
2,000
6,800
33,920
1,280
June 7
38,800
9,200
14,800
62 , 800
9,200
" 14
2,200
3,000
800
6,000
1,400
" 21
1,000
500
1,500
100
" 28
300
200
200
700
100
Bosmina longirostris has been frequently reported in the plankton
of European lakes. Apstein ('96) finds it perennial in Plonersee
with larger numbers in June-September and a maximum in July.
No pulse-like recurrence is noted, parthenogenesis prevails, and
males and ephippia are rare. His results, save in the matter of
pulses, are thus in general accord with ours. Stingelin ('97) notes
230
great seasonal polymorphism in B. cornuta near Basel. Zacharias
('97a and '98b) records it in the plankton of German carp ponds.
Stenroos ('97 and '98) finds it in waters of Finland and Karelia,
where the cornuta type is littoral, and a limnetic form, distinguished
by him as forma vernalis, is abundant in the plankton in May. Scour-
field ('98) finds it common in the waters of Epping Forest, where it
is perennial, males and ephippia appearing only in September-
November. According to Scott ('99) it appears at various seasons
in the lochs of Scotland in both the littoral and limnetic fauna.
Burckhardt ( 'OOa) gives an extensive revision of the genus Bosmina,
and includes in the B. longirostris group nine other so-called species,
among which are B. cornuta Jur. The species is "pelagic or hemi-
pelagic" in various Swiss lakes, though apparently not in num-
bers. The genus is there represented in the plankton princi-
pally by the B. coregoni group. Amberg ( '00) lists it from Katzen-
see, near Zurich, as a perennial planktont with large numbers in
May, August, and February, but gives no statistical data. Fuhr-
mann ('00) finds Bosmina perennial in Neuenburgersee, and B.
longirostris with a maximum in May. Marsson ('00) finds B.
" longirostris-cornuta " in lakes about Berlin throughout the year,
with larger numbers in some lakes during the warmer months and
in others in November-December. In Barlewitzersee, near Danzig,
Seligo ('00) reports B. cornuta as perennial, with maxima in June
and in October-November, the latter being the greater. Larger
numbers appear in summer than in winter. Cohn ('03), in waters
near Konigsberg, finds B. longirostris only sparingly present,
appearing in May-September with a maximum in July.
In European streams, also, B. longirostris is widely distributed.
Lauterborn ( '94) finds it abundant in the winter fauna of the Rhine.
He also states that it is not acyclic in the backwaters, where he has
found in three successive years both males and ephippia in May- June
and again in November. There is thus a suggestion of a vernal and
an autumnal pulse in these waters. Zimmer ( '99) finds it through-
out the whole year in the Oder. Schorler ( '00) reports it from the
Elbe at Dresden in May-October, with larger numbers in May- June
and September, while Fric and Vavra ('01) find it in the same
stream near Podiebrad. They state that B. cornuta is found in
great numbers in 1 m.- surf ace in summer months, and B. longiros-
tris sparingly in the littoral fauna. Steuer ('01) finds B, ' 'longirostris-
231
cornuta" m the backwaters of the Danube at Vienna in April-January.
It exhibits a distinct seasonal polymorphism, with a large winter
form and a smaller summer one. Data as to relative numbers
during the year are not given. Skorikow ('02), in reviewing the
investigations on the plankton of Russian waters, reports B. cornuta
from the summer plankton of several streams, but expresses doubts
as to whether "sie als autopotamische Planktonorganismen anzu-
sehen sind oder nicht." Meissner ('03) finds B. cornuta generally
in the Volga and its adjacent waters in the summer plankton, with
largest numbers in August; and Zykoff ('03) reports it in small
numbers from the same stream in May- July. It is not listed by
Volk ('03) in the Elbe at Hamburg.
B. longirostris occurs generally in American waters, though
apparently, often in small numbers. Thus Forbes ('82 and '90)
reports it in the plankton of Lake Michigan and Lake Superior, and
it appears generally in lists of Cladocera from many widely separated
smaller bodies of water in this country. Birge ('95 and '97) finds
only a few Bosmina (species not stated) in Lake Mendota, but
Marsh ('97) reports it (species not given) as perennial in Green
Lake, with a maximum in November. His records have also a
suggestion of an earlier pulse, in June, in which month there is a
sudden rise from a previous minimum.
This partial survey of the literature of the records of Bosmina
in the plankton shows its wide distribution, suggests the probability
of great variation, necessitating caution in the description of new
species in this genus, and indicates a wide diversity in its seasonal
career even in waters with somewhat closely similar environmental
conditions.
Ceriodaphnia me gaps Sars was found singly but once July 25,
1896, at 80.
Ceriodaphnia reticulata Jurine was found in the plankton occa-
sionally, and always in small numbers, in April-September. All
occurrences appear at temperatures above 66, and the earliest is
on April 1 7 , and the latest is September 2 1 . Females with summer
eggs were found in June-September.
Ceriodaphnia rotunda Straus was recorded in 1894-1895, but not'
thereafter. Its identification is somewhat questionable, and if
correct, this is apparently the first record of this species in North
American waters, unless it should appear that C. alabamensis
232
Herrick or C. acanthinus Ross, which appear to resemble C. rotunda
in some particulars, should be included here as forms or synonyms.
The genus is sadly in need of revision.
The forms referred to C. rotunda were found in August, 1894, and
July-August, 1895, 16,536 per m. 3 appearing in the plankton on
July 18 of the latter year.
Ceriodaphnia scitula Herrick. Average number, 1,539. This
species is closely related to the European C. quadrangula O. F. Mull.,
if, indeed, it is not identical with it. It is not impossible that it is
the form imperfectly described by Say ('18) as Daphnia angulata.
In the absence of a critical monograph of the genus I use the name
applied in current American literature to this form.
This is the most abundant species of the genus in our waters,
outnumbering all others by over sixfold in the totals of our records.
It is also one of the most important members of the Entomostraca
in the channel plankton (total of all records, 156,119), being ex-
ceeded in numbers only by Moina micrura (1,121,808), Bosmina
longirostris (381,598), Daphnia cucullata (237,444), and D. hyalina
(231,746).
It occurs in all months of the year except January and February,
but in larger numbers and in more of the collections in May-Septem-
ber. Thus less than 6 per cent, (reduced to 2 per cent, if one col-
lection in the warm autumn of 1897 is omitted) of the individuals
and only 20 of the 79 occurrences are found outside of the May-
September period. Ceriodaphnia scitula is accordingly a summer
planktont in channel waters. It is found in each year, though in
varying numbers according to hydrographic and other conditions.
Thus in 1898 the vernal pulse in June attains the unsurpassed
amplitude of 55,800 per m. 3 , but declines in a fortnight and makes
no recovery during the disturbed hydrographic conditions of the
summer. In 1897, on the other hand, our records were too meager
to delineate fully the vernal pulse, and in the stable conditions of
the summer and autumn the species continued in numbers whose
totals exceed those of 1898 by 81 -fold. Similarly in 1896 the more
gradual changes in levels which attended the floods of that year
permitted a considerable development of Ceriodaphnia throughout
the summer. Stable hydrographic conditions thus conduce to
increase in Ceriodaphnia. The relations which I have shown to
exist between Bosmina and movement in river levels (see table on
233
page 228) exist also in the case of Ceriodaphnia and in much the
same form.
The relation of temperature to Ceriodaphnia is evident in its
seasonal distribution. It does not advance rapidly in its vernal
increase until after the water warms to 70, and drops suddenly in
numbers when the autumnal decline passes this point. Moreover,
seasonal variations in temperature are accompanied by correspond-
ing shiftings of the pulses of Ceriodaphnia. Thus in 1898 the water
did not reach 70 until about May 20, reaching 73 on May 24, and
the vernal pulse of Ceriodaphnia began at once its rise to the maxi-
mum of June 7. In 1896 spring was early, 72 being recorded in
surface waters on April 24, and we find a vernal pulse rising to a
maximum on May 8. So also in 1897, when high temperatures
continued into the autumn, the decline passing 71 on October 5,
instead of in the first half of September as in other years, we find
the pulses of Ceriodaphnia extending into October with unusual
amplitude, reaching 5,200 per m. 3 October 5, while the highest
record in this month, or later, in other years was 280 per m. 3 Tem-
perature rather than season is thus the dominant factor in the
seasonal curve of occurrence of Ceriodaphnia.
The form of this seasonal curve is typically that of a series of
recurrent pulses of varying magnitude tending to reach the maxi-
mum height in the vernal pulse of May- June, attaining often lower
levels in July and rising again in August-September, and falling to
a minimum, or even to disappearance, in October. These later
pulses do not appear in the disturbed hydrographic conditions of
1898 (Table I.), but are clearly delineated in the summer records
of other years, especially in the stable conditions of 1897, where
well-defined pulses appear in July, August, September, and October,
at intervals of approximately four weeks, culminating July 14,
August 10, September 14, and October 5. Their maxima attain
respectively 5,600, 2,720, 6,000, and 5,200 perm. 3 , and the pulses are
delimited in each case by minima of less than 500 per m. 3 They
tend to coincide with those of other Entomostraca and to approach
those of the Rotifer a.
The Ceriodaphnia population in channel waters is almost ex-
clusively made up of parthenogenetic females. Males were not
recorded at any time, though females with ephippial eggs appeared
after the October pulse of 1897 and the vernal one of 1898.
234
Ceriodaphnia scitula appears but once in the records of European
plankton, Scourfield ( '98) finding it in the waters of Epping Forest
in September. The closely related C. quadrangula as well as the
other species have been frequently recorded by European investi-
gators both in the littoral and the limnetic fauna, but they appear
to be less generally found there than the other dominant Cladocera
of our waters.
It does not appear in the plankton of our Great Lakes (Forbes '82
and '90, Birge '95), or in that of Lake Mendota (Birge '95 and '97),
or .Green Lake (Marsh '97), but Herrick ('84) reports it as the most
abundant species in Minnesota, and Fordyce ('00) finds it in
Nebraska in shallow waters. A revision of the genus is needed
before the seasonal distribution of the various species can be worked
out on a basis that will make satisfactory discussions of the literature
possible.
Chydorus sphcericus O. F. Mull. Average number, 422, of which
26 are egg-bearing females, and 6 are immature, the remainder, 390,
being females in which the ova were not prominent.
The identification of species of Chydorus is attended by consider-
able uncertainty. Comparison with named specimens from Europe
supplied by Prof. G. O. Sars, leaves no doubt that C. sph&ricus is
common in our waters, and it is apparently the dominant species.
It is probable that several other species, as, for example, C. globosus
Baird and C. c&latus Schoedler, occur sparingly in our waters and
have been included with C. sphczricus in my enumerations. The
difficulties which attend the attempt to assign every individual to
one of the several species of Chydorus can be appreciated only by
one who makes the effort. The problem of their specific validity
should be solved by a statistical analysis of the range of varia-
tion.
The seasonal distribution of Chydorus sphcericus in channel
waters is in its general outlines very characteristic and well defined.
The following table, which gives the average number of Chydorus
per m. 3 for each month of our collections, shows clearly that it is a
vernal planktont, and that there is a slight tendency toward an
autumnal pulse in September, when vernal temperatures return.
The number for November (222) would probably be considerably
reduced if more than one collection had been taken in that month
in 1896. Omitting this year, the average for November falls to
235
78, and a secondary, hiemal rise becomes apparent in December.
This December pulse of Chydorus is one of the elements in the
upward movement of production in this month (see Part I.), and
fuller data may serve to connect it fully with the September-October
pulse, especially in more stable conditions. Both of these autumnal-
hiemal movements have less than one tenth of the development that
the vernal pulse exhibits.
The number and percentage of occurrences also confirm the
conclusions drawn from the numbers per m. 3 Percentages run
higher in the spring, in March-May, and in September-October
and in December, and lower in June- August, November, and
January-February. Chydorus occurred in all March collections,
and in only one third of the August collections.
The analysis of the data in this table indicates the presence of
Chydorus in the plankton practically throughout the whole year in
the whole seasonal range in temperatures, with the larger develop-
ments following shortly after the thermograph passes the yearly
mean (57 average of monthly means of surface waters) in vernal
rise and autumnal decline, the maximum development in April-May
SEASONAL DISTRIBUTION OF CHYDORUS. AVERAGE NUMBER PER M. 3
Year
Jan.
Feb.
March
April
May
June
1 804.
234
1895
11
2 044
1896
304
167
1,682
10,271
5,701
448
1897
20
540
320
32 800
900
1898
160
256
300
3,364
356
1 S99
36
65
193
Average
167
53
668
3,235
13,955
388
No of occurrences
9
6
15
9
9
10
Percentage of occur-
rences
75
40
100
82
90
72
236
SEASONAL DISTRIBUTION OF CHYDORUS. AVERAGE NUMBER PER M. 3 continued.
Year
July
Aug.
Sept.
Oct.
Nov.
Dec.
1894
95
461
100
16
56
1895
91
103
164
38
203
448
1896
64
104
78
160
800
277
1897
213
40
407
650
64
115
1898
50
30
60
28
172
1899
Average
103
49
228
202
222
214
No. of occurrences
11
7
13
12
10
14
Percentage of occur-
rences
61
33
81
71
63
82
occurring in average temperatures, for these months, of 60.5 and
68.3, while the minor autumnal development appears in September-
October at 74.2 and 57.6 respectively, and the December pulse, if
indeed it be a separate and independent pulse, is at the low tempera-
ture of 35.2. The December movement may be simply the result
of the more stable conditions which attend the appearance of the
ice-sheet on the approach of winter.
An analysis of the course of the seasonal distribution of Chydorus
in channel waters, as given in Table I. and in statistics of other
years, indicates the following seasonal regimen. In January-Feb-
ruary, at minimum temperatures, the occurrences are irregular
(75 and 40 per cent.) and the numbers small (average, 167 and 53
per m. 3 ), while in March, with rising temperatures, occurrences are
more numerous (100 per cent.) and numbers rise to 668 per m. 3 In
April-May a high percentage of occurrences (82 and 90 per cent.)
continues, and they mount rapidly to the maximum record of the
year, which in our statistics varies from 4,088 in 1895 to 32,800 in
1897. This vernal pulse reaches its maximum in our records on
April 29 in 1895, at 64, and in 1896 on the same day, at 70; on
237
May 25 in 1897, at 66. 3; and on May 24, in 1895, at 73. From this
maximum the pulse declines abruptly in a fortnight to a midsummer
minimum during maximum temperatures, which continues until
September. During this period the numbers are small, rarely rising
above 400 per m. 3 (average, 388, 103, and 49), and the occurrences
are also less numerous (72, 61, and 33 per cent.). With the decline
of temperatures which begins in September the percentage of occur-
rences mounts to 81, and the average per m. 3 to 228, and remains
near this level during the remainder of the year.
An analysis of the full statistical data, of which the records for
1898 are fairly typical, confirms the conclusions drawn from these
averages. Chydorus in channel waters is monocyclic, with a well-
defined vernal pulse in March- June which includes 95 per cent, of
the total annual Chydorus population. There are suggestions of an
autumnal pulse, but the data are not sufficient to delimit it. There
is no satisfactory evidence that there are recurrent cycles or pulses
at briefer intervals during the year.
The dominating effect of temperature as a regulating factor in
delimiting the seasonal distribution of Chydorus is very evident.
This, in addition to its appearance in the annual curve of occurrences,
is also exhibited most clearly in a comparison of the vernal pulses
in the two years of fullest representation in our records, 1896 and
1898. The following table gives the data of dates, temperatures
of surface waters, and numbers of Chydorus.
From these facts it appears that the late spring of 1898 delayed
the vernal pulse of Chydorus, and that the early spring of 1896
accelerated it in that year so that their apices (April 29 and May 24)
are four weeks removed from each other in seasonal location. In
both years the rapid rise in the pulse appears after 60 is passed, the
culmination occurs at about 70, and the decline, in temperatures
above 70.
Egg-bearing females were more abundant during the rise of the
pulse, and less numerous during its decline. Evidence of great
mortality during the decline of the pulses is to be found in great
increase in the relative numbers of empty carapaces. Thus, during
the decline of the vernal pulse in 1896 there were on the day of
culmination, April 29, 2,780 dead to 18,904 living, on May 1, 3,570
to 14,875, and on May 8, 1,578 to 6,706. From 14 to 24 per cent, of
the Chydorus population had thus recently perished. Parasitized
238
1896
1898
Date
Tempera-
ture
No. of
Chydorus
Date
Tempera-
ture
No. of
Chydorus
Mar. 17
42
256
Mar 15
46
440
" 24
40 7
610
" 22
51
480
" 30
48.1
6 405
" 29
49 5
240
Apr. 10
46 4
1 666
Apr 5
48 3
200
" 17
66 3
4 515
12
52
200
" 24
72
1 S 000
" 1
56
" 29
68
18 904
" 26
57
800
May 1
68 8
14 875
60
8
76
6 706
" 10
62
600
" 18
71 2
1 143
" 17
64
3 300 .
" 25
75 3
80
" 24
73
7 880
June 6
79
320
" 31
June 7
70
78
5,040
600
" 11
73
320
14
82 3
200
and fungused individuals were also noted in these periods of decline.
Males were recorded in September, December, and February.
Chydorus is not given as a constituent of the plankton of Nor-
wegian lakes by Huitfeldt-Kaas ( '98) or of Swiss lakes by Fuhrmann
('00), Amberg ('00), or Burckhardt ('00 and 'OOa). Its absence
from these cooler waters stands in sharp contrast with its abundance
in warm and shallow European lakes. It is reported as abundant
in Chroococcacea-rich lakes of North Germany by Apstein ('96),
where it is acyclic, with larger development in April-October, and
maximum in August or in May- June. According to Weismann
(79) Chydorus in some waters is polycyclic. It is also reported by
Zacharias ('97a and '98b) from the pond fauna of Trachenberg and
many other German localities, where it forms '"em notorisches
Mitglied des Teichplanktons." He also lists it ('98b)- from some
German streams. Marsson ( '00) found it in some waters near Berlin
in April- August, noting a great abundance in one instance in May.
Seligo ('00) gives a few statistical data indicating the occurrence
of Chydorus in the plankton of Hintersee near Danzig in April-
December, with a maximum in August and a secondary one in
October. It was, however, sparingly present in adjacent waters.
Cohn ('03) finds a like irregularity in its occurrence in waters near
Konigsberg.
Stenroos ('97) finds it to be one of the most abundant Entomos-
traca in the waters of northern Russia and ( '98) a littoral and bottom
species near Helsingfors. Scourfield ('98) finds it to be one of the
most abundant Cladocera in the waters of Epping Forest, occurring
from March to December, with maxima of sexual reproduction in
April and November. Scott ('99) reports it as abundant in the
littoral fauna of Scottish waters, but rare in tow-net collections in
open water.
It also occurs in the potamoplankton of European streams,
Zacharias ('98b) listing it from a few minor streams, but without
seasonal, statistical, or temperature data. It was not separately
listed by Skorikow ('97) in the summer plankton of the Udy at
Charkow, or by Lauterborn ('94) in the winter plankton of the
Rhine. Zimmer ( '99) found it from February to July in the Oder,
and Schorler ( '00) finds it abundant in the plankton of the Elbe in
April. Steuer ('01) finds it at all seasons in the backwaters of the
Danube at Vienna, and in the plankton from March to November
"oft in grossern Mengen," but gives no statistics of its seasonal
distribution. Fric and Vavra {'01) find it in the channel and
backwaters of the Elbe near Podiebrad, but more' abundant in the
littoral fauna, though no quantitative or statistical data of its
occurrence are given. Zykoff ('03) reports it as present in the
plankton of the Volga at all times in small numbers, and suggests
a predominance in May- July. Meissner ( '03) also reports it for the
Volga, but states that it is predominantly a member of the littoral
fauna though present in the plankton of the stream in restricted
numbers. No statistical data are given by him. Volk ('03) reports
it in the Elbe at Hamburg, but without any details.
This species is reported generally from American waters. Forbes
C90) reports it in the summer plankton of Lakes Superior and
240
Michigamme in small numbers, and ( '93) in that of the Alpine waters
of Wyoming and Montana, where it is, however, more abundant in
smaller pools. Birge ('94) finds it generally distributed in collec-
tions, including plankton, in Lake St. Clair and ('97) a member of
the plankton of Lake Mendota, where its abundance is dependent
on the supply of Anab&na. Its maximum only a single well-defined
one occurring in each year was found in July-October. Birge
regards it as an accidental member of the limnetic fauna, maintained
there as long as suitable food is present. Its mode of occurrence
does not, however, differ from that of typical plankton organisms,
which would doubtless likewise disappear from the plankton if their
food should be lacking.
It is noteworthy in this connection that it was only sparingly
present in the channel of the Illinois in the midsummer-autumn
plankton, when as, for example, in 1897 Anab&na and its allies
were abundant. It seems "not improbable that temperature even
more than food is an important factor in controlling its seasonal
and local distribution. It is unquestionably a member of the
plankton in our waters, though also abundant here, as elsewhere, in
the littoral fauna. In our locality in channel plankton it shows
distinctly seasonal limitations which suggest the operation of tem-
perature rather than food. Its occurrence in large numbers in
Wisconsin lakes in midsummer and its absence in the Illinois at
that time may also be correlated in part with the contrasted tem-
perature conditions in the two localities. Its occurrence in our
littoral fauna may also in part be due to the lower temperatures
consequent upon spring-fed areas and the shade of aquatic vegeta-
tion. Chydorus is one of those organisms capable of both the littoral
and limnetic habit under suitable conditions of food and temperature.
In our waters, at least, and, as it seems from the data of distribu-
tion, elsewhere, temperature, rather than food directly, appears
to be the factor controlling the occurrence of Chydorus in the
plankton.
Daphnia cucullata G. O. Sars. Average number, 181. In 1897,
very much greater, 5,483 per m. 3
For the reasons given by Burckhardt ('00) I use Sars's name
cucullata rather than jardinei of Richard to designate those forms
of the subgenus Hyalodaphnia in our plankton. In channel waters
this species varies considerably, but not to the extent that it does
241
where its numbers are greater. The forms known as apicata
Kurz and kahlbergiensis Schoed. appear in small numbers in some
years.
This species appears in our collections in April-December only,
with the exception of one occurrence in January and two in March.
Its occurrences and numbers vary greatly in different years. In
1894-95 its numbers were small and occurrences scattering, it being
most abundant in November-December. In 1896 there was a
large vernal development in April- June, and a series of diminishing
pulses in July-September. In 1897 no vernal development appeared
in our scattered collections, but in the stable conditions of late
summer and autumn occurred the largest development recorded in
any year, with a maximum record of 72,760 per m. 3 on October 5. In
1898 there was a small vernal development (3,400) in May- June
and a still smaller one (600) in October. A well-defined seasonal
routine is thus not demonstrable from our data, though the fact
that both the percentage of occurrences and the numbers are highest
in May- June and September-October suggests a tendency toward
vernal and autumnal pulses separated by a period of less develop-
ment in midsummer and of autumnal decline followed by a period
of almost complete extinction in midwinter.
The statistics of the D. cucullata population in all years in which
weekly collections were made, exhibit very clearly the phenomenon
of recurrent pulses of 3 to 5 weeks ' duration, with maxima of varying
amplitude and minima of less than 400 per m. 3 in all cases but those
which mark the September pulse of 1897. There are in 1896 pulses
culminating April 24 (2,544 per m. 3 ), May 8 (11,965), June 11
(12,000), July 18 (1,040), August 8 (800), and September 16 (507).
In 1897, vernal records are incomplete. Pulses appear July 14 (800),
August 17 (1,680), September 14 (57,000), October 5 (72,760), and
November 15 (2,040). These pulses coincide exactly or approxi-
mately with those of the other Entomostraca which exhibit the same
phenomenon, and approximate also those of the Rotifera. A typical
pulse, that of October, 1897, is shown in the following table. It
is a noticeable fact that the proportion of immature forms is often
greater at and after the period of maximum development than at
other times, as appears in the table.
The relations of temperature to the development of D. cucullata
in channel waters appear in the fact that all occurrences in excess of
242
Date
Females
Females
with eggs
Young
Total
Percentage
of young
Sept 27
160
320
640
1 120
57
." 29
7,520
4,000
12,800
24,320
52
Oct. 5
3,560
10,800
58 400
72 760
82
12
1 600
7 600
9 200
83
" 19
560
840
4 440
5 840
76
600 per m. 3 are found after the temperatures pass 70, with the[single
exception of the decline of the October pulse and the rise of the
November pulse to 2,040 per m. 3 at 47, following the high tempera-
tures in the late autumn and stable conditions of 1897. From the
depression in numbers during the period of maximum heat in mid-
summer and the occurrence of the major vernal and autumnal pulses
before and after its reign it appears that the temperature optimum
for D. cucullata in channel waters lies below this level, that is,
below 80.
D. cucullata is evidently very easily affected by the changes in
hydrographic conditions. Thus, in July-December, 1897 and
1898, the total movement in river levels was 12.4 and 31.4 ft.,
respectively, while the total cucullata population for these months
was 186,420 and 1,140 164-fold greater in the more stable year.
D. cucullata thus exhibits the maximum sensitiveness among the
Entomostraca to these environmental factors.
The D. cucullata population in the plankton consists almost
entirely of parthenogenetic females and young. The immature
stages form about 60 per cent, and the egg-bearing females 16 per
cent, of the total individuals. Dead, parasitized, or fungused indi-
viduals were found at times of the maxima or shortly thereafter,
but never in very large numbers. Males were found once in
December, 1896, and ephippial females also but once, on October
19, 1897, during the decline of the maximum pulse in our records.
Daphnia cucullata var. apicata Kurz, in well-developed condi-
tion, was found in relatively small numbers during the vernal pulses
of 1895 and 1896 and the autumnal pulse of the former year.
243
Incipient stages of this variety appeared also at other times. Burck-
hardt ('OOa) does not even concede varietal standing to apicata,
regarding it merely as a form of seasonal or local value. Its occur-
rence in our plankton when reproduction and growth are most
active suggests that it may have a growth value, and be in some
way correlated with the factors involved in its cyclic production.
Daphnia cucullata var. kahlbergiensis Schoed. appears but once
in our records in the plankton of June 11, 1896.
The D. cucullata group is a cosmopolitan constituent of the
fresh-w r ater plankton, appearing frequently in the records of Euro-
pean plankton. Apstein ( '96) finds it in lakes in northern Germany
in April-October with maximum numbers in July. The seasonal
limits thus resemble those in the Illinois, but the maximum falls
at the time of our midsummer decline. Temperatures in these
German lakes (16.3 C.) do not, however, reach the high levels
attained in our waters in midsummer. Stenroos ('98) records it in
several varieties in the plankton of Nurmijarvi See, the helmeted
varieties being found in midsummer. Zacharias records it from
the plankton of German ponds. Scourfield ('98) finds it in small
numbers in Epping Forest interruptedly in April-November, a
season coinciding with that in the Illinois. Burckhardt ( '00) finds
it represented by five different "forms" in Mauensee in the June
plankton. Marsson ('00) finds representatives of Hyalodaphnia
(species not given) in the April- June plankton near Berlin. Am-
berg ( '00) states that this species appears in April, increasing to a
maximum in July- August, and disappears again at the end of
November, a seasonal course similar in limits but not in maximum
to that in the Illinois. His data are too scattered to trace the course
of production with completeness. Seligo ('00), in waters near
Danzig, finds the species present in June- January, with maxima
in June- July and October. In the period of maximum summer
temperatures (16-21 C.) the numbers decline as in this period in
the Illinois. In Seligo 's infrequent (two to three weeks' interval)
data there are suggestions of minor recurrent pulses in other months.
Cohn ('03) finds in Lowentin a Daphnia w r hich he calls D. galeata
with vars. kahlbergiensis and cederstromii, and includes all three in
his enumeration. His investigation covers the months of May-
September, throughout which these forms appear, rising in a series
of recurrent maxima on June 26, August 4, and September 2 and 29.
(17)
244
Cohn seems not to have called attention to these clearly denned
recurrent pulses.
In European streams D. cuciillata also forms an important part
of the plankton. Lauterborn ('93) states that, with its varieties
kahlbergiensis and cederstromii, it appears abundantly in the plank-
ton of the Rhine in summer, but is not found in it in winter. Zimmer
( '99) states that D. kahlbergiensis was found constantly in the plank-
ton of the Oder in July-September, and Schorler ( '00) also finds it
in the Elbe at Dresden in May- August, with larger numbers in
June and August. Steuer ('01) reports it, in small numbers only,
in August in the backwaters of the Danube at Vienna. Fric and
Vavra ('01) reportD. kahlbergiensis as rare in the Elbe. Sowinski
( '88) finds it in several varieties in plankton of the Dnieper and its
tributaries, Rossinski ('92) finds it in the summer plankton of the
Moskwa, and Zernow ('01) in the June-July plankton of the
Schoschma and Wjatka. Meissner ('02 and '03) finds it in several
varieties in the May- August plankton of the Volga.
D. cucullata in some of its various forms or varieties appears to
be widely distributed in American waters. It was reported by
Forbes ('82), as D. retrocurva, from the plankton of Lake Michigan,
and also ('90) from Lake Superior and adjacent waters. Birge
('91 and '94) also finds it abundantly in Wisconsin waters and
in Lake St. Clair. Herrick ('84) and Ross ('97) report it from Min-
nesota and Iowa. Careful studies of its seasonal and vertical
distribution in Wisconsin waters have been made by Marsh ('97)
in Green Lake, and by Birge (95 and '97) in Lake Mendota. In
Green Lake D. kahlbergiensis is reduced to a minimum or even
extinction in December- April, rises in a late vernal maximum in
June- July, falls again to a lower level in August-September, and
then rises to a second and sometimes higher autumnal pulse in
October. In its main outlines this conforms to the seasonal course
of the cucullata form in our channel plankton. Our vernal maxi-
mum appears somewhat earlier, as a result probably of an earlier
warming up of the water. According to Birge ('97) this species is
more definitely periodic in its occurrence in Lake Mendota, being
confined entirely to July-December. Here also the largest numbers
are found in October, and the individuals gather in lower levels as
temperatures decline.
245
Daphnia hyalina Leydig. Average number, 417. In channel
waters this species has appeared in but two years, in 1895 in
April-July, attaining on June 19 a maximum of 166,208 per m. 3 , of
which 150,626 were immature. The collections were too infrequent
in these months to trace the course of this vernal pulse. D. hyalina
did not reappear until the spring of 1898, on May 24, in a single
vernal pulse culminating at 1 1,600 per m. 3 on June 7, and disappear-
ing a fortnight later. Its occurrences with one exception were all
at temperatures above 70. There is no apparent - cause for its
absence in later months or in other years. Males and ephippial
eggs were not found.
Daphnia hyalina is an exceedingly variable species, and a large
number of forms have been described which belong to the hyaUna
group. Burckhardt ('00), for example, recognizes 26 such forms
as varieties of this cosmopolitan planktont. This variability and
the difficulties attending the resulting synonymy cause any discus-
sion of the species in other waters to be attended by much uncer-
tainty. I shall therefore not attempt to distinguish in my dis-
cussion between the various varieties included by Burckhardt in
the hyalina group.
In lakes of northern Germany, Apstein ( '96) finds that D. hyalina
is essentially a winter planktont with a seasonal range of September-
July, and with maximum numbers in November- January. The
maximum thus appears there at the time of complete extinction in
our waters. Stenroos ('97) records it (as D. galeata) in the summer
plankton of Karelia, Huitfeldt-Kaas ('98) finds it in Norwegian
lakes in July and September in considerable numbers, and Scour-
field 's careful studies ( '98) of its seasonal occurrence in waters of
Epping Forest reveal an interrupted distribution in April-Novem-
ber. vScott ( '99) finds it in numbers in Scottish lochs in the plankton
examined at long intervals in March-January. Fuhrmann ('00)
reports it as perennial in Neuenbergersee, with a maximum in June
followed by a midsummer minimum. Burckhardt ('OOa) finds
great diversity in different Swiss lakes and in different years in the
relative numbers present. His intervals of collection were too great
to detect any pulse-like movement in the production, and it may be
that the diversity is due in part to the incompleteness of his records.
He concludes that D. hyalina is at a minimum in March-May,
increases in numbers slowly (with a preponderance of young indi-
246
viduals) in May-October to a maximum in November-January,
which is followed by a rapid decline (with preponderance of adults)
to the minimum. His results agree with those of Apstein ( '96) in
the main rather than with ours in the Illinois. Seligo ('00) finds
D. hyalina in Hintersee, though it is apparently absent from the
adjacent Barlewitzersee. In the former lake it appears in May,
rising to the year's maximum early in June, continuing throughout
the summer in diminished numbers, and disappearing in October.
In his infrequent records there are suggestions of several recurrent
minor pulses during the summer. Cohn ('03) reports D. galeata
regarded by Burckhardt ( 'OOa) as a form of D. hyalina from the
region of Konigsberg, but refers it rather to the cucullata group. I
shall therefore consider his results only in connection with D.
cucullata.
D. hyalina appears but rarely in the records of European potamo-
plankton. Steuer ('01) reports it, in small numbers only, in May
from the 'backwaters of the Danube at Vienna. Fric and Vavra
('01) state that D. microcephala regarded by Burckhardt ('OOa)
as a form of D. hyalina is abundant in the plankton at a depth of
0-1 m. in April-November in the Elbe and its backwaters at
Podiebrad. It is also reported by Zykoff ('00 and '03) in the late
vernal (June- July) plankton of the Volga at Saratoff, and by Meissner
('02 and '03) in the same stream in May- June. The examination
of the plankton of the Volga made by these authors is far less
extensive than that made of the Illinois River plankton, but as far
as it goes it indicates a similar distribution of D. hyalina in the two
streams. Volk ('03) reports it from the Elbe at Hamburg without
data.
The species appears to be widely distributed in American waters,
being reported, in some of its various varieties or synonyms, especially
from lakes and ponds. Smith ( '74) finds it in the plankton of Lake
Superior, Forbes ('82) in that of Lake Michigan, and Birge ('94)
in Lake St. Clair. It was also found in the Illinois by Forbes ('78)
and in the backwaters of the Ohio River by Herrick ('84), who
reports it also from Minnesota waters. Birge ('91) finds it in lakes
about Madison, Wis., and Fordyce ('00) in deep pools in western
Nebraska. The only investigation of its seasonal distribution in
American waters is that of Birge ('95 and '97) in Lake Mendota,
where it forms about 3 per cent, of all the Crustacea. It is perennial
247
in this lake but exhibits great differences in its seasonal course from
year to year. The vernal development in May-June (the only one
in our channel plankton) is relatively large in each year, but is
sometimes exceeded by an autumnal one in October. A midsummer
minimum sometimes appears between these pulses, and a winter
minimum in December- April is always present.
From the data here reviewed it seems probable that the very
limited seasonal distribution and irregular annual recurrence of
D. hyalina in our channel plankton is in a measure indicated in
streams elsewhere, and may have its cause in the instability of the
fluviatile environment as compared with the lacustrine, where the
species evidently finds its environmental optimum.
Diaphanosoma brachyurum (Lievin). Average number, 479, of
which 154 are females, 49 females with eggs, and 276 immature.
This species in our waters is monocyclic, with sharply defined
seasonal distribution. With the exception of two records of young
individuals in March- April, 1895 (and the identification of these
individuals is questionable), all our records of occurrence in
1894-1899 fall between May 25 and October 19, the first vernal
records appearing at temperatures of 55.8 to 72.3, and the last
autumnal at 52.5 to 65. The one pulse in each year except in
1894, when none was recorded falls in a period of 3-6 weeks in
July-September, the first record above 2,000 per m. 3 appearing
July 26, and the latest (with one exception, 2,175 on September 27,
1895) on September 7. The pulse varies in duration in different
years from 3 to 6 weeks, and attains a maximum on dates ranging
from July 26 to August 31, and varying in amplitude from 8,580
to 19,602 per m. 3 An analysis of the distribution of 61 recorded
occurrences in channel plankton shows that of these only 13, or 21
per cent., occur outside of July-September, and that the records
outside of the seven weeks of the pulse include less than 12 per cent,
of the total individuals.
A comparison of the seasonal curve of distribution with the
annual thermograph reveals the fact that the pulse occurs toward
the close of the period of maximum summer heat, and in every case
at a temperature of 78 or above, and that the decline of the pulse
often begins with declining temperatures, and is always accom-
plished during the autumnal decline. The effect of summer heat
pulses upon the Diaphanosoma curve is strongly suggested by the
248
data of the appended table, which gives the statistics of temperature,
river level, and Diaphanosoma population during the periods of
maximum development in 1895-1898. All these data except those
of Diaphanosoma are shown graphically in Part I., Plates IX.-XII.
The data for Diaphanosoma are less complete than the others, since
all of the collections were not counted.
In 1895 the Diaphanosoma pulse culminates at 19,602 on August
21, following immediately upon a heat pulse which culminates
August 15 at 85.3. The decline of the pulse occurs with a decline
of temperature to 72 on September 7. The declines, both of
Diaphanosoma and temperature, are hastened after September 3 by
1895
1896
Date
River
gage
Temp.
No. of
Diaph-
anosoma
Date
River
gage
Temp.
No. of
Diaph-
anosoma
July 2
" 1 n
5.15
4nn
80.8
7r> e
40
cnn
" 1 Q
2 en ic\
July 23
5.20
80
424
" 23
4.20
80
120
" 29
5.38
75.5
240
" 28
6.40
82
7,440
Aug. 1
4.20
78.5
1,088
Aug. 3
8.50
80.3
160
8
2.63
79
988
8
8.40
86
14,260
" 12
2.40
84.8
9,801
" 15
7.40
82
2,240
" 21
2.08
81.5
19,662
" 21
7.10
79
880
" 29
2.58
80
7,950
" 26
' Of)
6.50
77.5
7/1 1
600
Sept. 5
5.70
74
189
Sept. 16
4.10
73.5
663
" 12
3.90
79
1,053
" 30
4.30
58
80
" 20
7 ?n
7Q
468
" 27
3.23
73
2,175
1
249
1897
1898
Date
River
gage
Temp.
No. of
Diaph-
anosoma
Date
River
gage
Temp.
No. of
Diaph-
anosoma
July 14
6.30
79
160
July 12
7.00
78
60
" 21
5.20
81.1
960
" 19
4.70
84
40
" 30
4.60
84
4,720
" 26
2.90
89
8,580
Aug. 10
2.30
80.8
7,600
Aug. 2
2.70
78.3
6,960
" 17
1.90
79
7,120
9
3.20
83
360
" 24
1.80
77.5
5,120
" 16
3.70
77
60
" 31
1.80
80
1 1 , 000
" 23
4.20
82
1,020
* 3O
7 on
09 cr
7 con
Sept. 7
1.80
80
7,600
Sept. 6
4.70
79
240
" 14
2.00
83
1,500
" 13
4.20
62.5
1,800
" 21
2.00
71
240
" 20
4.20
73
960
" 27
4.90
73
400
the rise in river levels. Prior to that date hydrographic changes
are slight. With falling levels and higher temperatures after
September 7 there is a slight recovery in Diaphanosoma from 189
per m. 3 on the 5th, to 1,053 on the 12th.
In 1896 a well-defined heat pulse culminates August 10 at 86.5;
and Diaphanosoma, on August 8 at 14,260, with an abrupt depression
from 7,440, on July 28, to 160, on August 3, in flood waters. The
decline of this pulse from the maximum on the 8th to 440 on the 29th
is attended by a uniform decline in temperatures from 86 to 74.3 in
fairly stable hydrographic conditions, that is, declining river levels.
In 1897 there are two well-defined summer heat pulses, one
culminating August 3 at 89, and the other September 14 at 83,
separated by a depression to 77.5 on August 24. The crest of the
Diaphanosoma pulse likewise has two apices, the first culminating
at 7,600 on August 10, followed, during the decline in temperatures,
250
by a fall to 5,120 on the 24th, and, in the rising temperatures which
then ensue, by a recovery to a second maximum of 11,000 on the
31st. Diaphanosoma then declines though temperatures continue
to rise. These fluctuations all take place in comparatively stable
hydrographic conditions. There is a suggestion in the records of
this year that rising temperatures in midsummer conditions tend
to accelerate, and falling temperatures to depress, development of
the Diaphanosoma pulse, and also that after the pulse has continued
for some time (six weeks in this instance) rise in temperature ceases
to be effective. The autumnal decline in Diaphanosoma may
therefore not always of necessity be due to temperature decline alone.
In 1898 there are also two midsummer heat pulses, culminating
on July 26 at 89, and August 30 at 82.5, separated by a depression
which reaches 77 on August 16. The depression to 78.3 on August
2, with the consequent appearance of a third summit at 83 on
August 9, is due mainly to the fact that the temperature was taken
at 9:15 a. m., while all the others were in the late afternoon. The
seasonal curve of Diaphanosoma shows likewise two apices, the first
at 8,580 on July 26, and the second at 2,520 on August 30, separated
by a depression to 60 per m. 3 on August 16, when temperatures are
lowest. In this year the flood of the middle of August doubtless
plays a large part in depressing alike the thermograph and the
seasonal curve of Diaphanosoma, but in the light of the evidence
from 1897 in stable hydrographic conditions the direct influence
of temperature is also possible in this instance.
Diaphanosoma is thus a late summer planktont w r hich in develop-
ment is very responsive to changes in temperature. It appears in
the plankton in small numbers shortly after the establishment of
summer temperatures in May- June, but does not begin its maximum
development until maximum summer temperatures have existed
for six to eight weeks, and is apparently incited to this by a summer
heat pulse.
Males were recorded on July 18 and August 1, and ephippial
females on August 1 and September 5. Dead individuals were
most numerous during or subsequent to the maximum of the pulse.
This species is reported by Apstein ('96) in the plankton of
Dobersdorfersee, where it is also monocyclic, first appearing in
May, and attaining its maximum in September, when the males
first appear. In contrast with conditions in our waters the maxima
251
appear after the period of maximum summer heat. Zacharias ('97a)
reports it from German carp ponds in July, and Stenroos ('97) lists
it as a littoral species in midsummer in northern Russia. Scott ( '99)
finds it rarely in lakes of Scotland in August, and then only in the
plankton, though many shore collections were examined. Burck-
hardt ( '00) reports it from the smaller and shallower Swiss lakes in
isolated records ranging from May to November, and regards its
absence from the deeper lakes as due to the low temperatures which
at all seasons would surround its winter eggs, which sink to the low r er
levels. In Vierwaldstattersee ('OOa) he finds this species in the
plankton only in September-November, and then more abundantly
near shore than in the middle of the lake. In Alpnachersee the
period of occurrence extends from June to November with a maxi-
mum in July. Fuhrmann ('00) gives the seasonal distribution in
Neuenburgersee as extending from May to November, with a maxi-
mum in September. Marsson ('00) finds a seasonal distribution
from July to October in small lakes near Berlin. Seligo ( '00) finds
in Hintersee, near Danzig, a seasonal distribution in 1898 extending
from June 6 to October 18, with a maximum of 225,000 under
1 sq. m., depth, 24 m. (?) on August 9. Fric and Vavra ( '01 ) state
that this species is very abundant in summer months in the plank-
ton of the backwaters of the Elbe, especially in levels at depths
of 0-1 meter. Cohn ('03), on the other hand, finds in waters near
Konigsberg that Diaphanosoma is present in greatest abundance
in depths of 20-30 meters. It occurs in summer months, with large
numbers in July-September and a maximum in August-September.
It was not found in shallow waters.
As a constituent of the potamoplankton Diaphanosoma has been
reported by Schorler ('00) in the Elbe at Dresden as abundant in
June-September. Steuer ('01) finds it in the backwaters of the
Danube at Vienna in June-September, with a maximum in August,
but never in great numbers. Meissner ('03) reports it sparingly
from the Volga in July.
In American waters Diaphanosoma is widely distributed. Forbes
('90) found it abundant below surface levels in Lake Michigamme
in August. Birge ( '94) reports it in the plankton of western Lake
Erie but not in that of Lake St. Clair in September. In Lake
Mendota, Wis., he ('95 and '97) has worked out its seasonal and
vertical distribution with a fulness and care not equaled by any
252
European author previously quoted. Our results in Illinois waters
are in striking confirmation of his conclusions. He finds the first
scattering individuals in the plankton late in May, but numbers do
not rise until late in July or early in August, increasing rapidly
through August or even into September, then declining rapidly, and
disappearing entirely before November 1 . The active period is thus
at a time when a considerable part of the lake is at or above 68. In
our waters these temperature limits are 78 or above, but the sea-
sonal distribution is almost identical with that in Lake Mendota. He
finds it more abundant in the upper strata, 0-2 meters, than in the
deeper ones just the opposite of Cohn's ('03) results. Marsh
('97) has also determined its seasonal and vertical distribution in
Green Lake, Wis., with considerable care. Occurrences from the
last of October to the last of June are very few, and maximum
numbers appear from the middle of August to the middle of Septem-
ber, when surface waters have a temperature of 65-80. It occurs
in all depths (0-40 m.), but 70 to 80 per cent, of the individuals
were taken within 10 to 15 m.of the surface, the upper 5 meters being
more densely populated by night than. by day and in September-
October than in August.
Diaphanosoma is a typical planktont, with strong antennae, and
an active swimmer. Examination of the literature indicates its
wide distribution in the plankton of lakes and streams, and its very
marked seasonal limitation to seasons of higher temperature. It is
thus, as Birge ('97) has stated, markedly stenothermous. The
divergent conclusions concerning its limnetic habit and its vertical
distribution will doubtless be found to rest in some cases upon
insufficient data, and in others, upon its reactions to varying condi-
tions of light and temperature.
Eurycercus lamellatus O. F. Mull. This species occurred spar-
ingly and irregularly in the winter plankton at minimum tempera-
tures from November 30 to March 28. It is evidently adventitious.
Ilyocryptus spinifer Herrick. Average number, 4. This species
occurred sparingly and irregularly in the plankton during the
warmer months. The earliest record was on July 23, and the latest
October 11 at 65. This species is evidently adventitious in the
plankton. I have doubtfully referred our examples to Herrick 's
species /. spinifer, for the reasons given by Herrick and Turner ('95),
rather than to /. longiremis, to which Birge ('91) would refer our
253
American form described by Herrick as /. spinifer. A larger amount
of material exhibiting a fuller range of variation may, however, serve
to connect the two.
Leptodora hyalina Lilljeborg. Average number, 3. This species
occurred in small numbers and somewhat irregularly in our collec-
tions of channel plankton in summer months. Our earliest record
was June 28 ; and the latest, August 30. It is our largest crustacean
planktont and a fairly active swimmer, and was often taken in our
tow-nets, which had a larger mouth and coarser mesh (No. 12) than
our plankton net. I took this species in great numbers in the upper
meter of water at midday in May- June in Lake Meredosia with a
seine of No. 000 silk. It may be that it is less abundant in the
channel than in the backwaters, and the small number in the plank-
ton collections from the channel may also be accounted for in part
by the escape of Leptodora from the small orifice (10 cm.) of the
plankton net, or to its negative rheotropism when stimulated by the
currents of the plankton pump.
Macroihrix laticornis Jurine was found in the plankton in May
at 64-73, adventitious in flood waters.
Moina micrura Kurz. Average number, 261 per m. 3 In 1897
it was much more abundant, averaging 5,106 in the more stable
conditions of that year.
This is the most abundant of all our Cladocera, appearing in
great numbers in periods of stable low water during maximum
temperatures. It is exceedingly irregular in the extent of its devel-
opment in different years, the average numbers per m. 3 in 1894-1898
being respectively 21,844, 22,842, 188, 5,106, and 261. After mak-
ing allowances for the irregularity in the number and distribution
of the collections in the several years, it still remains apparent that
Moina is very uneven in its distribution.
The seasonal distribution of Moina in channel plankton is con-
fined to July-September with the exception of 9 occurrences in
small mmibers in the last days of June and the early part of October.
The earliest record is June 19, in 1895, w r hen the very large number
of 329,448 per m. 3 were found, a degree of development which
implies a previous period of multiplication. The first records in
subsequent years were all later than this date in June or early in
July. After several recurrent pulses, each of 3 to 5 weeks ' duration,
the numbers decline to a very low level, and the species disappears
254
from the plankton in September-October. In 1898 (Table I.) the
last record was made October 11 the latest in any year with the
exception of an isolated record October 26, 1897. Moina micrura
is thus distinctly a summer planktont.
It appears in the plankton only after maximum summer tem-
peratures of approximately 80 have been reached, and decreases
rapidly as soon as the autumnal decline passes this point, and soon
thereafter vanishes from the plankton. Its optimum temperature
in channel waters is thus near 80.
The relation which hydrographic conditions bear to the ap-
pearance of Moina in channel plankton appears upon a comparison
of the Moina population and the movement in river levels in differ-
ent years, as shown in the following table.
MOINA AND HYDROGRAPHIC CHANGES.
June
July
August
September
October
S
^
s
^
B
^
8
^
s
d
d
a
S
S
d
V
S
B
0)
**
B
O
E
^j
E
E
O
E
o
E
B
s
B
B
Year
. E
>
O
E
O
**
. E
>
o
. E
o
G
g S3
E
^ OJ
E
&
E
Z CD
p.
I
a
!*
&
, i
p,
,,
P,
^_i
M
+3
M
5
H
S
M
a
bo
$
>
?>
O
t>
^
o
j>
o
^
^
h
^
H
^
H
^
1894
192
3.4
40,415
2.1
129,880
2.6
3,677
4.7
3.1
1895
329,448
2.7
91,318
7.3
2,597
3.5
87
8.8
10
2.7
1896
3.4
152
7.8
1,220
4.3
3.7
4.6
1897
6.3
1,373
5.2
1,280
2.6
70,040
0.6
605
0.6
1898
75
4.0
660
7.4
1,496
7.5
770
6.2
40
3.9
While the correlation is not proportionate between the extent of
movement in levels and the Moina per m. 3 , it is still very evident
that in years of continued and more stable low water Moina is
found in much greater numbers, as appears on a comparison of 1897
and 1898. It is also confined largely to the more stable part of the
year, appearing in 1895 in June- July in large numbers, but falling
off when the minor floods of August-September occur, while in 1897
the large numbers are found in the stable levels of August.
255
The cause of this limitation of Moina to periods of low levels in
maximum temperatures appears to lie in the food relations of the
species. Moina abounds in waters approaching stagnation. The
slackened current, increased sewage contamination, and excessive
growth of the smaller algas and chlorophyll-bearing flagellates at
such seasons in the channel of the Illinois furnish an environment
favorable to the great increase in Moina, such as was recorded in
the low water of July- August, 1894, of June- July, 1895, and of
September, 1897, exceeding in each instance that of any other
species of Entomostraca in the plankton. The relatively smaller
numbers of Moina at the same seasons in the less contaminated
backwaters lends additional support to the view that these condi-
tions approaching stagnation are in a measure responsible for its
unusual development in channel plankton.
Of the total Moina population, over 65 per cent, are young or
immature, 7 per cent, are egg-bearing females, embryos are often
freed from the parent on application of the preserving fluid, 11 per
cent, are males, and the remainder, females without eggs. Males
appeared with the maximum or decline of the major pulse for the
year in 1894 (August), 1895 (July), 1897 (July and September), and
1898 (September), but ephippial females were recorded only in
June-July, 1895.
The seasonal distribution of Moina conforms to the type of a
series of recurrent pulses wherever the numbers are considerable
and the collections sufficiently frequent to delineate their courses.
Even in the small numbers of 1898 (Table I.) there are suggestions
of such pulses.
Moina micrura seems to be a species characteristic of the pota-
moplankton. It is not mentioned as a constituent of the plankton
or littoral fauna by any of the various investigators quoted else-
where in this paper who deal with lakes or ponds in Europe or North
America; nor does it appear as a frequent constituent of the
potamoplankton elsewhere. Skorikow ('02), indeed, makes the
statement, " Bemerkenswert ist fur die Fliisse vollstandiges Fehlen
der Gattung Moina." This, however, is hardly the case, for
Sowinski ("88) finds it in the plankton of the Teterew, a tributary
of the Dnieper, and Fric and Vavra ('01) report it from the Elbe
in 0-1 m. strata in July-September, males appearing in the latter
month. Meissner ( '02 and '03) also finds it in the Volga at Saratoff,
256
where it " appears almost constantly in the plankton." His investi-
gations, however, appear to cover only the months of May-August.
Maximum numbers appeared in July, and considerable differences
were noted in two successive years.
I find no previous record of the occurrence of Moina nticrura in
American waters.
Pleuroxus denticulatus Birge. Average number, 5. Occurs in
small numbers and irregularly during the autumn and spring months
during declining or rising temperatures. The earliest autumnal
record is November 2, and the latest, December 15; the earliest
vernal is March 8, and the latest is May 31. Egg-bearing females
appear in the earlier occurrences in each season. It is evidently
adventitious.
Pleuroxus hamatus Birge was found once March 29, 1898.
Scapholeberis mucronata O. F. Mull, was recorded in small num-
bers in May and August-December through the seasonal range of
temperatures. It is apparently adventitious in channel plankton,
though not attending flood invasions.
Sida crystallina O. F. Mull, is rare in the summer plankton.
Simocephalus serrulatus Koch. Average number, 261. This
species appears irregularly in the plankton, generally in small
numbers and in isolated occurrences. An exception to this is found
in May- June, 1898 (Table I.), when it is found continuously May
10- June 14 in numbers which furnish 61 per cent, of the total for
all years. There is a slight preponderance of occurrences in May
and September, 12 of the 26 recorded appearing in these months.
Their irregular appearance in the plankton in general suggests that
they are adventitious from the littoral area, especially at times
of their maximum development there. The period of their occur-
rence in the channel plankton in 1898 was one of rising water, 10 to
14 feet above low-water mark a stage permitting free communica-
tion between the channel and large areas of slightly submerged
bottom-lands.
Simocephalus vetulus O. F. Mull, appeared irregularly and in
small numbers in the plankton in April- June (4 occurrences) and
September-December (5 occurrences). It is evidently adventitious
in the plankton, coming from the littoral area, though not confined
to flood waters.
257
i OSTRACODA.
The species of this order are in the main, during adult life,
limicolous forms found in the littoral or bottom ooze or amid the
decaying organic matter which accumulates in these regions. The
current, the movements of fish and other large aquatic organisms,
the action of waves along shore and in shoal regions, all tend to bring
these animals into the limnetic fauna. Their centers of distribution
are thus in littoral or bottom regions, and in the adult stage they
are almost wholly adventitious in the plankton of our waters. In
1898 the average number per m. 3 was 191, but in 1897, a more stable
year, only 97.
The seasonal distribution of their occurrences in the plankton
indicates a decided predominance in March-October, in which
months all but 6 of the 73 records were made. In these months
from 23 to 82 per cent, of the collections contained Ostracoda, while
in December-February only 8 to 20 per cent. The percentages in
April-September are all above 45 per cent., and the numbers per
m. 3 are also larger in this period (see Table I.). The tendency
toward a vernal increase is apparent in the records of each year in
much the form in which it occurs in 1898 (Table I.). The numbers
are always small at all seasons, not exceeding 1,600 per m. 3 even
in the vernal season.
The seasonal distribution is such that the greater part of the
occurrences and the greater number of individuals appear in the
plankton during the warm season, that is, above 50. Thus, in 1898
all but 4 of the 24 occurrences and 99.5 per cent, of the indivi-
duals appear after the vernal rise passes 50 and before the
autumnal decline reaches that point. The Ostracoda are plank-
tonts of the warmer season.
It is significant that the Ostracoda in our plankton collections
are largely young or immature individuals. In 1898, 'for example,
74 per cent, of individuals observed were not adult, and most of
these appeared in April-June. Their occurrence in the plankton
can not be traced to the action of flood waters. It thus seems
probable that the young Ostracoda may temporarily adopt more
of a limnetic habit than the adults.
No attempt was made to systematically identify the Ostracoda of
the plankton catches. The list of species and the notes thereon
258
which follow, are drawn in the main from Sharpe ('97), to whom I
am also indebted for assistance in identifications which I have
made. A few supplementary notes are based on my plankton
.records.
DISCUSSION OF SPECIES OF OSTRACODA.
Candona sigmoides Sharpe. is rare in shore collections below the
plankton station.
Candona reftexa Sharpe was taken but once in the river on
November 11.
Candona simpsoni Sharpe appears commonly in April-May, and
again, in smaller number, in October-November in shore collections
on the west side of the river at the plankton station. It is occa-
sionally adventitious in the plankton at these seasons.
Cypria exsculpta Fischer appears rarely in the channel plankton
and in shore collections in April-October.
Cypria ophthalmica Jurine is found frequently in the plankton
throughout the year, but more abundantly in May-September, and
especially in late summer and early autumn.
Cypria pustulosa Sharpe was taken rarely in channel plankton
in July and September.
Cypridopsis vidua O. F. Mull, was perennial in the plankton,
though present in greater numbers in May-October. It is the
commonest of the Ostracoda in the plankton, and it seems probable
that many, though not all, of the young and immature forms belong
to this species.
Limnicythere illinoisensis Sharpe was taken in the plankton in
March, August, and November in 1898, in two instances in flood
waters.
COPEPODA.
This is the most abundantly represented order of the Entomos-
traca in channel plankton. Though the species number but 12 to
the 25 Cladocera, the individuals among the Copepoda outnumber
the Cladocera over fivefold in the grand totals, the ratio varying in
individual years from twofold in 1894 to almost sevenfold in 1898.
The average number in 1898 was 40,608 per m. 3 ; in 1897, in
more stable conditions, 80,632; in 1896, a year of recurrent floods,
43,764 approximately the number in 1898; in 1895, a year of low
w r ater in spring, 116,264 the highest average of any year; and in
259
1894, 53,149. On June 19, 1895, the Copepoda attained a vernal
maximum of 1,022,476 per m. 3 more than twice the maximum
record for any other year.
The Copepoda occur in every collection examined, and throughout
the whole seasonal range in temperatures. As shown in Table I.,
the copepodan population during minimum temperatures in De-
cember-February is at a minimum, the number per m. 3 rising above
10,000 per m. 3 in but 6 instances in 44 collections in these months,
and falling below 1,000 in but 5. In March- April, as temperatures
rise, the numbers increase rapidly, especially after 50 is passed, to
a vernal maximum in the last days of April or early in May, usually
at the time of the vernal volumetric maximum or very shortly
thereafter. In fact, volumetric maxima are generally accompanied
by copepodan maxima culminating at the same time or a week
later, as in May, 1898, when the volumetric is on May 3 and the
copepodan on May 10.
Numbers continue to be large during the period of summer
heat, declining somewhat tardily with the autumnal decline in
temperatures. In midsummer in 1898 numbers fall below 20,000
in 9 instances in disturbed hydrographic conditions, but in all
previous years in April-September there are only 9 such records in
a total of 63. The decline to the winter minimum is usually com-
pleted in November, though in 1897, 20,000 is not permanently
passed until December 21, at 32.
The Copepoda are thus perennial in the plankton, and the fact
that they exhibit a larger winter population than the Cladocera is
due to the fact that a number of species, the Harpacticida, Cyclops
bicuspidatus, C. prasinus, C. serrulatus, and C. modestus appear to
be planktonts belonging to the colder part of the year. As a whole,
however, the Copepoda reach their greatest quantitative develop-
ment in the warmer part of the year, with a major pulse in April-
May and an occasional autumnal pulse, as in 1897, of equal or
greater proportions.
The whole course of the seasonal occurrence of the Copepoda as
revealed by collections at frequent intervals, exhibits the phenome-
non of recurrent pulses at intervals of 3 to 6 weeks, and more clearly
denned in stable conditions. Owing to their relatively smaller
numbers the adult Copepoda do not show the pulse phenomenon
( 18)
260
as clearly as the nauplii and immature forms. In 1898 the adults
form only 10 per cent, of the total.
The relation which hydrographic conditions bear to the
copepodan population may be inferred in part from the comparison
of years given above, and from the following table, in which are
given the average number of Copepoda per m. 3 and the total monthly
movement in river levels in July-December, 1897 and 1898.
July
August
September
1897
1898
1897
1898
1897
1898
Average Copepoda
per m. 3
81,543*
5.2
7,720
7.4
121,070
2.6
11,080
7.5
261,387
0.6
36,920
6.2
Total movement in
levels in ft
October
November
December
1897
1898
1897
1898
1897
1898
Average Copepoda
per m. 3
128,093
0.6
28,285
3.9
49,240
2.2
10,692
2.6
15,740
0.5
7,908
2.4
Total movement in
levels, in ft. ...
With a total movement of 1 1.7 ft. in July-December in 1897 and
nearly three times as much (30 ft.) in 1898, we find copepodan
population falling off to less than one sixth that of the more stable
year.
Of the total Copepoda in our records for 1894-1899, 78 per cent,
are nauplii of Cyclops and Diaptomus, 13 per cent, are immature
Cyclops, and the remaining 9 per cent, are HarpacticidcB, Diapto-
mus, and adult Cyclops. Of the twelve forms, Cyclops viridis var.
insectus is the most important quantitatively, and includes one
fourth of the total adult copepodan population, exceeding the
next in importance, C. viridis var. brevispinosus, by over threefold.
261
The following forms are of numerical importance in the order
named : C. bicuspidatus , young Diaptomus, Cyclops edax, Diaptomus
sicilbides, D. pallidus, Canthocamptus spp., and Cyclops albidus.
Cyclops prasinus, C. modestus, C. phaleratus, and C. serrulatus are
also found, but in such small numbers as to be of no quantita-
tive consequence.
DISCUSSION OF SPECIES OF COPEPODA.
Argulus sp. A small and apparently young argulid was found
in the plankton on August 10, 1897. Members of this genus are
abundant upon Amia calva and both species of Lepisosteus, all very
common fish in channel waters.
Canthocamptus spp., including C. illinoisensis Forbes. Average
number, 78. Canthocamptus was found in the plankton in every
month of the year but June. The percentage of collections contain-
ing Canthocamptus is greatest (44 to 63 per cent.) in March-May and
November, and the numbers per m. 3 are highest in March-May,
when females, females with eggs, and nauplii all occur in their
maximum numbers. All records of totals in excess of 400 fall in
this vernal period with the single exception of one collection in
August, 1897. The largest number, 3,058 per m. 3 , was found
April 29, 1896.
Canthocamptus occurs throughout the whole seasonal range in
temperatures, with smallest numbers and least regularity during
maximum summer heat in June- August. It is thus a planktont
of the colder rather than the warmer part of the year.
The relations which hydrographic conditions bear to the occur-
rence of Canthocamptus in the plankton may be inferred from the
fact that of the 48 records in 1894-1899, 24 were made in rising
flood waters, 14 in falling flood stages within a few days after the
culmination of the rise, and but 10 in stable conditions or in declining
levels when flood waters of recent origin did not fill the channel.
From these facts it seems probable that Canthocamptus is in the
main adventitious in the plankton from its normal habitat in the
slime at the bottom and margins of the river and its backwaters.
Over 88 per cent, of the total Canthocamptus recorded in the
plankton consists of nauplii. It may be that as is the case with the
young Ostracoda they enter the area of the plankton more readily
than the adults. Adults were found in the plankton only in
262
November-May; females with eggs, only in February- April ; and
a female with attached spermatophore, in March. Nauplii appear
in greatest numbers in April-May, attaining 2,862 perm. 3 April 24,
1896, but they rarely rise* above 400 per m. 3 outside of this vernal
period, and are found only in very small numbers in December-
March. It appears from our data that the breeding season is prin-
cipally in April-May.
Cyclops albidus Jurine. Average number, 113; in 1897, 136; in
1896, 33; and in 1894, but 10. A discussion of the variation and
synonymy of this species has been published by E. B. Forbes ('97).
The species is numerically least important of the dominant members
of the genus in our plankton. It was recorded in all months but
December and February, but its season is practically confined to
April-October, the only exceptions being three records in small
numbers in January, March, and November, and two of larger
numbers (300 and 200) in the higher temperatures of the delayed
autumn of 1897. There is a tendency toward a summer minimum
in June- July, with pulses of greater amplitude in April-May and
again in August-October. In these months the percentage of
collections containing C. albidus is highest, being respectively 55,
50, 38, 56, and 53 per cent., and these are the only months in which
the numbers per m. 3 rise above 600. The highest numbers recorded,
2,862 and 2,400, occurred respectively on April 24, 1896, and
October 5, 1897.
Although C. albidus is found in the extremes of temperatures,
it shows a decided increase after temperatures pass 60 in the
vernal rise, and falls off immediately after the autumnal decline
passes this point. With high temperatures continued into October,
in 1897 we find it continuing in larger numbers. On the other
hand, during maximum summer heat (about 80) numbers, as a
rule, fall below 300 per m. 3 The temperature optimum thus appears
to be in the neighborhood of 70. The three greatest pulses re-
corded, occur respectively on April 24, 1896, at 72; on April 26,
1898, at 57-, and on October 5, 1897, at 71.
The numbers are too small to exhibit very clearly the phenome-
non of recurrent pulses, though the vernal and autumnal pulses are
usually well defined, and in the stable conditions of 1897, August,
September, October, and November pulses may be traced.
263
Hydrographic conditions appear to affect C. albidus as they do
other Entomostraca. In July-December, 1897, in stable low water
the C. albidus population exceeds by over threefold that of these
months in 1898.
Of the totals of all records in 1894-1899, 74 per cent, are fe-
males, 4 per cent, with eggs and 70 per cent, without, and the
remaining *26 per cent, are males. Immature forms and nauplii
were not distinguished from those of other species. Egg-bearing
females were recorded only in May and August-October, at times
of maximum pulses. Over 82 per cent, of the males were found
in August-October a period of declining temperatures and decreas-
ing food supply.
This is a widely distributed species, though it seems generally
to be present in relatively small numbers in the plankton. It occurs
in many European lakes. Stenroos ( '98) finds that it is the most
abundant species of Cyclops in Nurmijarvi See, occurring in both
the plankton and littoral fauna throughout the summer. Scourfield
('98) finds it common in the waters of Epping Forest, where it is
perennial in ponds and small lakes ; and Burckhardt ( '00) also finds
it in the smaller lakes of Switzerland.
It appears to be more generally reported from European streams.
Thus, Schorler ( '00) finds it to be rare in the plankton of the Elbe
at Dresden. in May; and Fric and Vavra ('01), perennial in the
littoral fauna of the same stream at Podiebrad, while Volk ('03)
reports it in the plankton at four of seven localities examined at
Hamburg. Meissner ('02 and '03) finds it in May-August in the
Volga at Saratoff, where it is abundant in the littoral zone or among
vegetation and in quiet backwaters.
Under a variety of synonyms this common and variable species
has been reported from many American waters by Herrick ('84)
and others. It was described by Professor S. A. Forbes ('90) as
C. gyrinus, from the plankton of Lake Superior. With the exception
of Marsh's record ('95) from Lake St. Clair, it does not elsewhere
appear to have been found in the plankton of the Great Lakes.
Marsh ('93 and '95) finds it generally in the plankton of smaller
bodies of water in Wisconsin and Michigan, and E. B. Forbes ('97)
reports it as generally distributed in American waters of a permanent
character. Brewer ('98) reports it (as C. signatus) in the vernal
plankton of deep pools near Lincoln, Neb. No statistical
264
data on its seasonal distribution are given by any of the authors
cited.
C. albidus appears thus to be adapted to both the littoral and
limnetic areas, but seems never to attain great numbers in the
latter.
Cyclops bicuspidatus Claus. Average number, 373; in 1897,
206; in 1896, 145; in 1895, 312; and in 1894, only 2. A full dis-
cussion of the variation and synonymy of this species has been
published by E. B. Forbes ('97).
This species shows sharply marked seasonal limitations. Every
one of the 68 records, with the exception of one of a single female
found September 30, falls within November-May, and all of the May
records were made in the delayed low temperatures of the spring of
1898. The general distribution of this species during this period
is indicated by the high percentage of collections in which it was
found, viz., 63, 71, 67, 73, 93, 53, and 40, respectively, for November-
May. The numbers per m. 3 are, however, high only in November
and April-May, reaching 8,000 in 1895 and 1898 in this vernal pulse,
and 3,560 in November, 1897, in the autumnal pulse. In Decem-
ber-March numbers do not rise above 500 per m. 3 save once in
December and on March 24-30, 1896. C. bicuspidatus is thus a
winter and early spring planktont in channel waters of the Illinois.
The temperature adaptations are exhibited by the fact that only
13 of the 68 occurrences are in temperatures above 50, only 5 above
60, and but 1 above 70 that of May 24, 1898, at 73. On the
other hand, the greater developments in numbers take place during
these higher temperatures of 50-70, the only rises above 1,000 per
m. 3 at temperatures below 50 being those of March 30 and April 10,
1896, at 48 and 46.4, and of November 15, 1897, at 47. Mini-
mum numbers thus prevail below 45, and the temperature opti-
mum in channel waters of the Illinois appears to lie near 60.
The seasonal routine in channel waters begins with the appear-
ance of small numbers about November 1 , with an occasional pulse
of some amplitude in that month followed by a continuance of small
numbers through the minimum temperatures of December-Feb-
ruary, and a rise with the temperatures in March to a maximum
vernal pulse toward the end of April or the first of May, and a
complete disappearance of adult individuals after temperatures pass
70 during May-October.
265
Stable hydrographic conditions appear to favor the increase in
C. bicuspidatus, as is seen in the large pulse of November 15, 1897
(3 ,560) , and the slight pulse (240) during declining levels in February,
1899.
The vernal development of 1898 (Table I.) is distinctly pulse-like,
and there are traces elsewhere of similar phenomena, but in general
the numbers of C. bicuspidatus are too small to exhibit clearly the
phenomenon of recurrent pulses.
Of the totals of all individuals recorded in 1894-1899 I find that
37 per cent, are males, 16 per cent, egg-bearing females, and 47 per
cent, females without eggs. Immature forms and nauplii were not
distinguished from those of other species. With the exception of
a few stragglers, the egg-bearing females were limited principally
to March-May. In exceptional cases the males greatly outnum-
bered the females, as on November 15, 1897, when the ratio was
2,820 to 680.
Though apparently widely distributed, this species does not
appear frequently among the planktonts reported from European
lakes. Scourfield ( '98) reports it as a common species in the waters
of Epping Forest throughout the year with the exception of a period
of absence or depression in July-August, and Scott ( '99) finds it in
shore collections made in various months of the year in Scottish
lakes, and more abundantly in the warmer months. It has been
reported in the potamoplankton in Europe only by Rossinski ('92)
from the Moskwa, by Zernow ('01) from the Schoschma, and by
Volk ('03) from but one of seven localities in the Elbe at Ham-
burg.
In American waters, on the other hand, C. bicuspidatus is more
abundant, and in the Great Lakes it forms a very important part
of the plankton. Forbes ('82) finds it (as C. thomasi] to be the
dominant Cyclops in the summer plankton of Lake Michigan and
('90) also abundant in that of Lake Superior. Marsh ('93 and '95)
finds it in the summer plankton of the Great Lakes, near Charlevoix,
in Lake St. Clair, the Detroit River, and Lake Erie, but only rarely
and in small numbers in the smaller bodies of water in Wisconsin
and Michigan. E. B. Forbes ('97) extends its recorded range to
Massachusetts and to the lakes and rivers of Wyoming, and states
that it is widely distributed in America and occurs in large ponds
and rivers. Brewer ( '98) reports it in the vernal plankton of deep
266
pools near Lincoln, Neb. None of the investigators quoted give
statistical data of the seasonal limitations of C. bicuspidatus.
The absence of this species from the summer plankton of the
Illinois River and its abundance in that of the Great Lakes is perhaps
explained by the temperature conditions. Surface waters in Lake
Michigan are reported by Ward ('96) to range from 62 to 67
August 11-29, while deeper waters at and below the thermocline
reach a minimum of 42. The warmest waters there (62-67) are
thus considerably cooler than the coolest in the waters examined
by us (which are usually above 70 and often above 80) during the
months in which C. bicuspidatus is not found in our plankton. That
its absence is not due to sewage contamination in low water which
usually prevails during the warmer months is shown by the prompt
reappearance of the species in the autumn; as, for example, in 1897,
when sewage was even more abundant than usual. It may be that
temperature is also one of the factors limiting its distribution
elsewhere.
Cyclops edax Forbes. Average number, 49; in 1897, 194; in
1896, 159; in 1895, 321 ; and in 1894, 187. This is the third species
of Cyclops in numerical importance in channel plankton of the
Illinois.
With the exception of a single record on November 2, 1897, all
occurrences of this species in channel plankton are confined to
April-October, and all but 9 of the 48 occurrences are in July-
October, and 32 of them in July- September the period of maxi-
mum summer heat. During these three months the percentage
of collections containing C. edax is highest (44 to 75 per cent.), and
they are the only months in which the C. edax population rises
above 1,200 per m. 3 in channel waters excepting a single instance on
October 5, 1897, in the high temperatures of that delayed autumn.
In other months the records are all below 800 and generally below
400 per m. 3 The highest number recorded was 3,600 on October
5, 1897.
The seasonal distribution, with maximum numbers in July-
September, exhibits a temperature adaptation on the part of C. edax
to maximum summer temperatures (70 to 80) in channel waters.
An examination of the records shows that only 13 of the 48 records
of this species fall in temperatures below 70, and these were all in
the months of April, May, September, October, and November, at
267
times when occurrences were scattering and numbers few; that
is, during the rise or decline of the species to or from the summer
maximum. Of the 13 records below 70, there were 5 between
60 and 70, 7 between 50 and 60, and but 1 below 50. Cyclops
edax in channel waters of the Illinois is thus stenothermic in narrow
limits near the maximum temperatures of the year.
The relation which hydrographic conditions bear to the seasonal
development of C. edax may be inferred from the fact that the
July-October population of this species in the disturbed waters of
1898 was only 35 per cent, of that in the more stable months of the
preceding year.
The occurrences of C. edax take the form of pulses, though less
distinctly recurrent and less clearly denned than in species present
in larger numbers. Such pulses appear in July, August, and
September, 1895, and in August and October, 1897. In 1898
(Table I.) the numbers present are too small to clearly indicate
recurrent pulses, though suggestions of the phenomenon appear in
the records. In general these pulses tend to coincide with those of
other Entomostraca.
Of the totals of all our records of C. edax in 1894-1899, 60 per
cent, are females without eggs ; 1 1 per cent., females with eggs ; and
29 per cent., males. Young and nauplii were not distinguished
from those of other species. Egg-bearing females were found in
April and in June-October, but in greatest numbers in July- August.
Males occur in June-November, with no marked predominance in
any period.
This species has not been separated from C. leuckarti by other
investigators of the plankton, though E. B. Forbes ('97), after a
careful comparison of American forms with C. leuckarti of Europe,
concludes that edax is specifically distinct, and that leuckarti also
occurs in American waters, though apparently not in numbers com-
parable with those in European waters. C. edax appears in a
measure to replace it in our plankton. He reports it as widely
distributed in American lakes and streams and in the plankton of
our Great Lakes.
Cyclops leuckarti Claus. A single dead specimen was recorded
in channel plankton August 26, 1898. E. B. Forbes ('97) records
it from the Fox and Sangamon (tributaries of the Illinois) , from the
Illinois and Mississippi rivers, and from Quiver, Flag, and Dogfish
268
lakes, backwaters of the Illinois at Havana. It is not, however, at
any time a factor of any importance in channel plankton of the
Illinois at Havana, being confined to the spring-fed lakes or those
shaded by vegetation, where regions of lower temperatures may be
found.
This is a widely distributed form in the plankton of European
waters. Stenroos ('98) finds it abundant in the plankton of Nur-
mijarvi See, Scourfield ( '98) reports it as common in the waters of
Epping Forest in February-October, and Scott ( '99) as rare in that
of Scottish lakes. Fuhrmann ( '00) states that it is always rare in
Neuenburger See except in April, and is absent in November-
December, while Burckhardt ('OOa) finds it to be perennial in
Vierwaldstatter See, with breeding season in May-September and
maximum in August or September.
It has been generally reported from European streams. Schorler
( '00) finds it in the Elbe at Dresden in May-October, with greatest
numbers in July-September, and Volk ('03) reports it from four of
seven localities in the same stream at Hamburg, though Fric and
Vavra ( '0 1 ) do not find it at Podiebrad. Zykoff ( '03 ) , Zernow ('01),
and Meissner ('02 and '03) find it in the plankton of Russian rivers.
The last author states that it occurs in both channel plankton and
littoral fauna among vegetation where breeding females abound
during the maximum in May. The young only appear in the chan-
nel plankton.
In American waters this species has often been held to include
C. edax, and the data here quoted from Birge and Marsh refer to the
combined species. Marsh ( '93 and '95) finds it generally distributed
in the lakes of Michigan and Wisconsin, and in the plankton of
lakes Erie, Michigan, and St. Clair. Birge ('97) finds it in the
summer plankton of Lake Mendota, where it is even more abundant
than C. viridis var. brevispinosus.
Cyclops modestus Herrick was recorded in channel plankton
only in November, December, and March, in small numbers and
isolated occurrences at temperatures of 41 and below. E. B.
Forbes ('97) states that this species lives in shallow, weedy water,
and has never been found in large numbers, though widely dis-
tributed. On account of its relative rarity it may have been over-
looked by me and have a wider seasonal distribution than my
scanty data indicate.
269
Cyclops phaleratus Koch was recorded in channel plankton only
in small numbers in November-December, 1897, at minimum tem-
peratures. E. B. Forbes ('97) states that it is a littoral form,
confined to marginal vegetation.
Cyclops prasinus Fischer. Average number, 2. This species
occurs sparingly and irregularly in September-March in channel
plankton, appearing in largest numbers in the early autumn of 1895
and most continuously in the winter of 1898-99. The numbers are
always small, never reaching 400 per m. 3 , and in 12 of the 17 records
falling below 100 per m. 3 The percentage of collections containing
C. prasinus in the totals rises above 20 per cent, only in December
(24 per cent.). The seasonal distribution in channel plankton
indicates a limitation to the colder part of the year, all records but
5 being below 40. Nevertheless, in September-October, 1895, the
species was recorded in 56-79. This fact and its relatively small
numbers generally, make it probable that inferences from our
scanty data concerning its seasonal distribution can not be con-
clusive.
Of the totals in all years, 86 per cent, are females without eggs,
6 per cent, females with eggs (found in February and November),
and 8 per cent, males.
E. B. Forbes ('97) finds the species widely distributed in
American waters from the Great Lakes to roadside pools. Marsh
('93 and '95) finds it (as C. fluviatilis} in the larger bodies of water
in Wisconsin and Michigan, and in lakes Erie, Michigan, and St.
Clair. In Green Lake he ('97) finds it to be the most abundant
species of Cyclops, and perennial, with maxima in September-
November. His statistical data exhibit somewhat irregular numbers
which contain suggestions of recurrent pulses such as appear in
our records of other species of Cyclops. Brewer ('98) finds the
species in the plankton of pools near Lincoln, Neb.
Cyclops serrulatus Fischer. Average number, 3. This species
was taken sparingly in channel plankton, exhibiting only isolated
occurrences in December, January, March, and May, in flood waters
at temperatures of 32-75. It is much more abundant in Spoon
River, where it is sometimes the dominant species of the genus,
appearing in May-September, and in small numbers in colder
months. It appears to be adventitious in channel plankton of the
Illinois River.
270
This widely distributed Cyclops appears but rarely in the records
of the plankton of European lakes, and then only in the smaller
ones. Stenroos ('98) reports it as abundant in the littoral zone of
Nurmijarvi See; and Scourfield ('98) finds it perennial and the
most abundant species of Cyclops in the waters of Epping Forest.
On the other hand it has been found generally in the plankton
of European streams. Zimmer ('99) finds it in the Oder, and
Schorler ( '00) states that it is abundant in April- June in the plank-
ton of the Elbe at Dresden ; Fric and Vavra ( '01) find it only in the
littoral fauna at Podiebrad; and Volk ('03) in the plankton in four
of seven localities in the Elbe at Hamburg. Sowinski ( '88) found
it in the plankton of the Dnieper, Rossinski ('92) in that of the
Moskwa, Zykoff ('00) in the summer plankton of the Volga, and
Zernow ('01) in the winter plankton of the Schoschma. Meissner
( '02 and '03) reports it in May- August as not abundant in the back-
waters and vegetation of the Volga at Saratoff.
In American waters Marsh ('93 and '95) finds it in smaller lakes
of Wisconsin and Michigan but not in the Great Lakes, and E. B.
Forbes ('97) states that it is one of the most common and widely
distributed species in American waters. It appears, however, not
to be quantitatively an important element in lake or river plankton.
Brewer ('98) finds it to be the most abundant vernal Cyclops in the
small bodies of water near Lincoln, Neb.
Cyclops viridis Jurine. A synonymy and a discussion of varia-
tions in this the dominant and most variable of all the Cyclops in
our channel plankton, has been given by E. B. Forbes ( '97). I have
grouped the individuals in our plankton under two varieties,
brevispinosus Herrick and insectus Forbes. The two varieties inter-
grade, and in my separation I have followed only a single character
readily visible without dissection or manipulation, namely, the
outer terminal spine of the stylet, which is short, broad, and lance-
shaped in brevispinosus, and more spine-like in insectus. Judging
from the results of this method of separation, it appears that this
lance-shaped spine is a character of the male in many instances,
though not found in all males or limited to this sex.
Cyclops viridis var. brevispinosus Herrick. Average number,
124; in 1897, 447; in 1896, 622; in 1895, 850; and in 1894,68. This
form occurred in all months but January, but predominantly from
the last days of April to the first week in October, the percentage
271
of collections containing brevispinosus in these months being 27,
80,62,67,48,75, and 5 9 per cent , respectively , while in other months
it does not rise above 20 per cent. The number of individuals is
also greater during the warmer season. No record between October
15 and April 20 exceeds 200 per m. 3 , while between April 20 and
October 15 the pulses often culminate at 3,000-5,000 per m. 3 , and
over 98 per cent, of the total individuals were recorded.
This variety appears throughout the whole seasonal range of
temperatures from summer's maximum to winter's minimum, but
predominantly during the warmer season. Only 15 of the 71
occurrences and 2 per cent, of the individuals w r ere recorded at
temperatures below 60. As soon as the vernal rise in temperatures
passes 50-60, the minimum numbers and scattered occurrences of
the winter months give way to a vernal pulse of considerable mag-
nitude in April-May, attaining 4,452 on April 25, 1895, and 4,960
on May 25, 1897, but only 2,600 on June 7, 1898. This is followed
by a period of depression in July, when the summits of the pulses
did not often surpass 1,000 per m. 3 In the late summer and autumn
of 1895 and 1897, and to a less extent in 1896 .and 1898, a second
period of maximum pulses appears, attaining 9,711 September 12,
1895, and 4,800 October 5, 1898. When temperatures decline in
September-October below 50, this variety falls at once to minimum
numbers.
The records of brevispinosus in channel plankton exhibit some-
what clearly the phenomenon of recurrent pulses whenever collec-
tions at brief intervals make it possible to delimit the pulses. Thus,
in 1895 there are pulses culminating in July, August, September, and
October; in 1896, in April, May, June, July, August, and September;
in 1898, in July, August, and October; but in 1898 (Table I.) the
numbers are too small to exhibit fully the phenomenon of recurrent
pulses.
The relation to hydrographic conditions may be inferred from
the fact that while in the stable conditions of July-October, 1897,
pulses culminated at 800-4,800 per m. 3 , in the same period in the
disturbed hydrographic conditions of 1898 no pulse rose above 200
per m. 3 , and the total of all records in those months is only 8 per
cent, of that in 1897. Evidently brevispinosus does not thrive in
flood waters.
272
The surprising fact derived from the examination of our records
of this variety of C. viridis, is that the individuals referred to it are
predominantly of the male sex. Out of a total of 74,308, 64,883, or
88 per cent., are males, 8,542, or 11 per cent., females without eggs,
and only 883, or one per cent., egg-bearing females. In so far as
these data go, they indicate that this so-called species, or even
variety, of C. viridis, in so far as it is based on the lance-like spine
of the stylet, is not well founded. This is, it seems, predominantly
a male character, though not exclusively so, since females, and even
egg-bearing females, are found which exhibit this structure.
C. viridis var. brevispinosus appears to be confined to American
waters. Marsh ('93 and '95) reports it from the larger lakes of
Wisconsin and Michigan, and from the Great Lakes, except Lake
Michigan. Birge ('95 and '97) finds that it is the most abundant
species of Cyclops (except in summer, when C. leuckarti abounds)
in Lake Mendota, and the only one reproducing under the ice. His
data exhibit a major pulse in May, and a second one, of less ampli-
tude, in October, with slight indications of recurrent minor pulses
in midsummer, obscured possibly by the massing of his data in
fortnightly averages. The seasonal distribution in Lake Mendota
is thus much like that in the Illinois River. Marsh ('97) finds the
maximum in Green Lake in June at 68-69, and only scattering
occurrences at other seasons. E. B. Forbes finds this variety
widely distributed in American waters, but never especially abun-
dant.
Cyclops viridis var. insectus Forbes. Average number, 539; in
1897, 2,115; in 1896, 949; in 1895, 2,966; and in 1894, 905. It is
thus more abundant by two- to threefold in the stable years of 1895
and 1897 than in the flood-swept years of 1896 and 1898.
This variety was found in every month of the year, though
predominantly in April-October, when the percentages of the
collections containing it were respectively 64, 100, 85, 100, 100, 87,
and 76 per cent. In November-March the percentages were only
44, 6, 17, 7, and 13. The numbers of individuals are very small,
however, from October 1 to April 20, excepting in the autumn of
1897, when, with the delayed high temperatures and the great
impetus given to plankton development in the stable conditions of
low water, the maximum pulse of all our records, 30,800 per m. 3 , was
reached on October 5, a pulse of 1,200 following in November. With
273
these exceptions no record exceeding 600 per m. 3 was made between
the dates named. Between April 20 and October 1 the minimum
records rarely fall below 600 per m. 3 , except in 1898, and the pulses
often culminate at 2,000-8,000. C. viridis var. insectus is thus a
planktont of the warmer season, and its seasonal distribution is
strikingly similar to that of the so-called var. brevispinosus.
This form occurs in our plankton throughout the whole seasonal
range in temperatures, but only in small numbers and irregularly
below 60. Only 21 per cent, of the collections containing
insectus were made at temperatures below 60, and these contained
less than 3 per cent, of the total individuals. With the exceptions
of the pulses culminating at 43 November 23, 1897, at 1,200 per
m. 3 , and at 57 April 26, 1898, at 4,160 per m. 3 , no development of
this species exceeding 600 per m. 3 occurs below 60. All pulses of
more than 3,000 per m. 3 , excepting only the April pulse of 1898,
occur at temperatures above 70. The species reaches its greatest
development in channel waters during the period of maximum
temperatures, 70-80.
The seasonal distribution of this form shows a few straggling
individuals in November-March during temperatures below 50,
and a meteoric rise to a vernal pulse in April-May as this tempera-
ture is passed and 60-70 arrives. This is followed by a series of
recurrent pulses, often of considerable amplitude, through Septem-
ber or until temperatures fall below 60, as in October, 1897. With
falling temperatures the drop in numbers to the winter minimum
is quickly accomplished. A comparison of the distribution in 1897
and in other years, shows a close correlation between the decline in
temperatures and the falling off in numbers of insectus.
The relations which hydrographic conditions bear to the develop-
ment of insectus in channel plankton may be inferred from the
hydrographs on Plates IX.-XII, Part I., and from the data sum-
marized in the following table, 1894 being omitted because of the
incompleteness of the seasonal representation.
In 1895 levels were low, unusually so in the spring, and the
flood-free intervals of the year were of more than the usual extent.
About 10 feet of the total movement in levels (51.9 ft.) is found in
the late December rise. If this is excluded, the total movement
falls to 42 feet, and the range in levels to 6.5 feet. Under conditions,
274
Year
Range in
levels, in ft.
Total
movement,
in ft.
Average height,
in ft., of stage
of river
Average number
of insectus
per m. 3
1895
12.2
51.9
3.61
2,966
1896
10.1
45.7
6.98
949
1897
14 3
44.8
6 90
2, 115
1898
15.5
67.2
8.02
539
then, of lowest levels, least range, and total movement, we find the
largest development (2,966) of insectus in channel plankton.
In 1896 the average river level is much higher, affording in-
creased current and more silt. A series of recurrent floods also
flush the channel, though the total movement and range in levels
within the limits of the year are not greatly increased. Neverthe-
less, the changes, which appear mainly below bank-height, affect
channel plankton profoundly, and the production of insectus falls
to 949 per m. 3 In 1897 the population rises to 2,115 per m. 3 , largely
as a result of the stable conditions of flood-free waters at low levels
and with slight current in the last half of the year. In 1898 the
total movement (67.2), range in levels (15.5), and average stage
(8.02) reach the extremes in the four years under comparison, and
the insectus population falls to the lowest level 539 per m. 3
A detailed comparison of the July-November period of the two
years follows.
Month
July
August
September
Year
1897
1898
1897
1898
1897
1898
Total movement
5.2
6.05
5,093
7.4
5.70
210
2.6
2.29
2,030
7.5
3.66
304
0.6
2.01
2,275
6.2
4.44
325
Average stage
Average number of C.
viridis var. insectus. . .
275
Month
October
November
Average
Year
1897
1898
1897
1898
1897
1898
Total movement
0.6
2.01
8,625
3.9
4.86
200
2.2
2.82
520
2.6
7.44
68
2.2
3.04
T.709
5.5
5.22
221
Average stage
Average number of C.
viridis var. insectus. . .
In 1898, with two and a half times the movement in. levels found
in 1897, the development of insectus attains less than 6 per cent, of
the numbers reached in the latter year.
The occurrences of insectus in channel plankton exhibit the
phenomenon of recurrent pulses during the season of its occurrence
in large numbers whenever collections are sufficiently frequent to
delimit the pulses. Thus, in 1895 there are. such pulses in July,
August, September, and October; in 1896, in April, June, July,
August, and September; in 1897, in July, August, September,
October, and November; and in 1898, in April, May, June, July,
August, and September, though of slight amplitude in the last three
months.
Some of the seeming gaps and irregularities in the series of pulses
of brevispinosus and insectus will be eliminated if the statistics of
the two forms are combined in a single series, a fact which lends
support to the view that the two forms belong to the same species,
and are parts of a common group of variable organisms.
Steuer ('01) concludes from his examination of the plankton
of the Danube at Vienna, based on 19 (?) collections in 15 months,
that Cyclops has usually two maxima and two minima in each year,
and that in the same body of water, owing to various meteorological
influences, the two maxima do not in any year fall near each other.
The more extensive data at my command show the limitations of
such a general conclusion. An examination of the records of indi-
vidual species of Cyclops and of the total Cydopida in our waters,
make it clear that the major pulses may follow each other at about
a monthly interval. For example, in 1897, the total Cyclopida
(19)
276
have their major occurrences in our records as follows, the pulses
appearing September 14 and October 5 :
July 30 8,080 Sept. 14 117,000
Aug. 10 49,360 Sept. 21 15,260
Aug. 17 17,120 Sept. 29 14,400
Aug. 24 20,320 Oct. 5 101,600
Aug. 31 67,200 Oct. 12 3,400
Sept. 7 107,200
Again, in 1896, the two major pulses of the year are on June 19
(928,984) and July 18 (563,815). Steuer's conclusion seems to be
founded upon insufficient data, and can not have general applica-
tion.
Of the total 240,830 individuals of C. viridis var. insectus in our
records in 1894-1899, 117,166, or 49 per cent., are males; 109,460,
or 45 per cent., females without eggs; and 14,204, or 6 per cent.,
females carrying egg-sacs. If the brevispinosus totals are included,
the percentages change to 42 per cent, of females of which 37 per
cent, and 5 per cent., respectively, are without and with egg-sacs
and 58 per cent., males. The apparently high proportion of males
may be due to the fact that in the enumeration more young females
than males were. included in the "young" Cyclops.
The egg-bearing females were generally more numerous in
April- July. No marked predominance in the proportion of males
appears at any season in our records.
Cyclops viridis does not appear extensively in the plankton
literature of European lakes. Stenroos ( '98) finds it not rare in the
littoral fauna of Nurmijarvi See. Scourfield ('98) reports it as
next in abundance to C. serrulatus in waters of Epping Forest, w r here
it is perennial. Scott ('99) finds it at all seasons in both littoral
and pelagic collections in Scottish lakes, and Amberg ('00) lists it
for Katzensee.
It appears but infrequently in the investigations of European
streams. Neither Schorler ('00) nor Fric and Vavra ('01) report
it from the Elbe, though Volk ( '03) lists it from six of seven localities
in this stream at Hamburg. Sowinski ( '88) finds it in the littoral
fauna of the Dnieper, and Zykoff ('03) in the summer plankton of
the Volga, though Meissner ( '03) states that it is never found in the
plankton of that stream at Saratoff, being confined to the littoral
277
zone and to vegetation. No statistical data concerning its seasonal
distribution are given by any of these authors, though Meissner
states that it reaches its maximum in May in the Volga.
In addition to the species of Cyclops here listed for the channel
plankton of the Illinois, E. B. Forbes ('97) records in May-Septem-
ber, 1896, C. varicans Sars as common, and C. fimbriatus var. poppei
Rehberg and C. bicolor Sars as rare.
Owing to the impossibility of separating with certainty the
nauplii and young of the various species of Cyclops they were all
recorded together under the head of " nauplii " and " young Cyclops."
The former includes also the nauplii of the two species of Diaptomus
occurring in our plankton.
Young Cyclops. Average number, 4,780; in 1897, 16,035; in
1896, 10,196; in 1895, 21,960; and in 1894, 5,960. With two ex-
ceptions in January and February they occur in every collection
examined. Numbers are, however, at a minimum in November-
March, only 9 instances of more than 1,500 per m. 3 appearing in our
records in this season. With the exception of two pulses in the
autumn of 1897, and two in this season in 1895, all pulses of an
amplitude exceeding 8,000 per m. 3 are confined to the interval
between April 20 and October 1, practically to temperatures above
70. They also exhibit relations to hydrographic conditions of the
same nature as those found in case of the adults of the various
species of Cyclops, and manifest likewise the phenomenon of re-
current pulses (Table I.). The totals of all young Cyclops in 1894
1899 are almost five times those of all adults of the genus. This
ratio gives an index of the extent of the decimation by enemies and
inimical factors of the environment which exists after the nauplius
stage has passed and before that of the adult is reached.
Nauplii of the Copepoda (excluding the Harpacticidce) . Average
number, 36,707; in 1897, 53,786; in 1896, 24,560; in 1895, 88,442;
and in 1894, 45,648. Nauplii were recorded in all collections ex-
amined with but two exceptions. As in the case of the adults and
young, the large numbers are, however, confined to the warmer
season between April 1 5 and October 1 . During the colder months
the pulses rarely rise above 20,000 per m. 3 , and those in excess of
35,000 during these months are with one exception confined to the
delayed high temperatures of the stable autumn of 1897. During
278
the warmer season, on the other hand, the pulses frequently attain
100,000 or over.
The maximum record of 928,984 was made in the stable low
water of June 19, 1895. All large developments thus lie at tem-
peratures above 70.
The nauplii bear much the same relation to hydrographic condi-
tions as that found in the adults; for example, in Cyclops viridis.
This is seen in the fact that in unstable years such as 1896 and 1898
the numbers are on the average only 28 and 68 per cent, of what
they were in the more stable conditions of 1895 and 1897, and the
average monthly population in July-December in the unstable
conditions of 1898 is only 18 per cent, of that in the same months
of the previous year.
The relative numbers of adult, young, and larval stages of the
Cydopida are given in the accompanying table.
Year
Nauplii
Young Cyclops
Adult Cyclops
No.
Ratio
No.
Ratio
No.
Ratio
1894
456,483
2,741,718
1,451,524
1,828,720
1,908,780
121,345
38
19
17
18
30
61
59,598
680,749
428,211
545,200
248,576
5,422
5
5
5
5
4
3
11,726
140,779
84,786
102,730
62,735
1
1
1
1
1
1895
1896
1897
1898
1899
Totals
8,508,570
21
1,967,756
5
404,749
1
The ratios between total adult and young, 1 to 5, are fairly
constant in the different years, falling to 1 to 3 in January-March,
1899, and to 1 to 4 in 1898, a year in which the colder part of the
year was most fully represented. This ratio probably represents
more truly the relationship of young and adult in the total yearly
production. The ratios of adults to nauplii in the several years
vary considerably from the totals of all years (1 to 21), rising to 1 to
279
61 in winter conditions of 1899 (January-March), and falling as low
as 1 to 17 in 1896. This was a year of recurrent floods, but its ratio
is in sharp contrast with that of 1898 (1 to 30), also a year of con-
siderable hydrographic disturbances during the summer. The adult
population was reduced during this year, and especially during the
summer floods, but the nauplii do not fall conspicuously below
those of other years. It would therefore seem that the deleterious
action of flood conditions operates more effectively upon the adult
and young than upon the nauplii. This fact may be due to the
relative absence of spines and hairs on the nauplii, structures which
gather silt and load down the larger forms in the flood, waters.
The greater number of young and adults in 1896 as compared
with 1898 may be due to the more gradual rise of the floods of the
former year (see PI. X. and XII., Pt. I.) jand the proportionally
greater amount of silt in the more sudden floods of the latter.
The ratios given in the table are of course subject to the error
arising from the uneven seasonal distribution of the collections in
some years, and to that arising from varying location of the collec-
tions on the pulses, especially on those of greatest amplitude. An ad-
ditional error arises from the leakage of the smaller nauplii through
the meshes of the silk net. I have found on experiment that they
will thus escape under pressure of a column of water only 3-4 cm. in
height. Their dimensions are such that the smaller individuals
can pass through the meshes of even the No. 20 silk. It seems
probable that ratios of nauplii to adults are actually greater than
our records indicate.
The relationship which the pulses of nauplii bear to those of the
adult Cydopidcz may be inferred from an examination of the data
of Table I. An analysis of the seasonal distribution of the total
young and adult Cydopidce and of the nauplii reveals the fact that
in all seasons in which collections at approximately weekly intervals
were made, their pulses coincide in a majority of cases in their
maxima, and when the coincidences do not occur the maximum of
the nauplius pulse appears in the collection of the week following
that of the young and adult Cydopidcs. This appears less constantly
and clearly in the disturbed hydrographic conditions of 1898 (Table
I.) than in the records of more stable years.
Apstein ('96) finds that nauplii of Copepoda are most abundant
w r hen eggs are most common, and that this bears no constant relation
280
to the abundance of adults. Our collections, extending over longer
periods and being at briefer intervals, indicate, however, that this
relation does exist. As above stated, the larvae are most abundant
at or shortly after the times of greatest abundance of adults that
is, the maxima of the recurrent pulses. Apstein also states that
reproduction is periodic and development rapid. Maximum
numbers are reported by him in May and September.
Cohn ( '03), on the other hand, maintains that the " innere Logik"
and his data show him that the nauplii reach their greatest numbers
just prior to the appearance of largest numbers of young and adult
Copepoda. His data are from 12 collections between May 1 and
October 1 , and favor his contention in 2 out of 3 cases (of maxima) ,
and both of these lie in collections at intervals of 15 to 16 days. In
the light of our data obtained at briefer intervals and the conclusions
therefrom that the pulses of larvae tend to coincide or follow at a
brief interval those of the adults, it becomes questionable whether
his data are sufficient for his conclusion. His logic also overlooks
the fact, apparently, that smaller numbers of larvce might lead to
coincident maxima of grown forms during a period of abundant
food, on which all pulses must be based, since the larval stage may
be at such times a brief one and the adult a relatively longer one, and
the cumulative effect of this relationship would make the conditions
shown in our data logically possible. Furthermore, Cohn used a
No. 12 silk in his plankton net, and this allows many nauplii to
escape, and probably accounts for the fact that the ratio of larvas
to grown forms in his figures is only 1.3 to 1, while in our records it
is 3.5 to 1. The discrepancy arising from this leakage may further
tend to weaken his data for his conclusions concerning the relations
of larvas and adults.
Steuer ('01) finds that the nauplii in the Danube at Vienna
reach maxima in June and in August, but his data are too scattered
to fully delineate their fluctuations. Two out of three of his max-
ima coincide with those of all Cyclops, and the third antedates it
(monthly intervals of collection), as in Cohn's data.
Diaptomus pallidus Herrick. Average number per m. 3 , 11; in
1897, 367; in 1896, 87; in 1895, 152; and in 1894, 146.
This species was recorded in all months of the year but February,
though in a larger percentage of the collections and in larger numbers
in July-December. Prior to this season the percentage does not
281
rise above 31 per cent., the occurrences are irregular, and the num-
bers are small. Thus in 1896 and 1898, years of numerous winter
and vernal collections, there were but 4 occurrences in each prior to
July 1 , and all but one of these was of numbers less than 100 per m. 3
Only 12 of the 72 occurrences and 8 per cent, of the total individuals
were recorded in the first and less stable half of the years. In
July-December numbers rise in feebly outlined pulses which attain
at the most 800-2,400 per m. 3 The percentage of collections con-
taining the species rises to 33-75 per cent., and in stable autumns
such as 1895 and 1897 the occurrences are but little interrupted. In
its seasonal distribution in channel waters it is thus largely confined
to the last and more stable half of the year.
Its relationship to hydrographic conditions here suggested also
appears in a comparison of the yearly averages given above. The
average numbers per m. 3 in 1896 and 1898, 87 and 11, are greatly
exceeded by those of 1895 (152) and 1897 (367). The total number
recorded in July-December in 1897 is 29 times that in 1898. This
well-defined predominance in stable seasons, which appears also in
the case of the closely related D. siciloides, exceeds that of the other
Entomostraca, and indicates a greater sensitiveness on the part of
these species to the deleterious effects of flood waters. The long
antennas and great development of the feathering of the caudal
stylets afford a large area for the attachment of the silt and debris
of flood waters, and accordingly facilitate the destruction or removal
of Diaptomus from the plankton more quickly than in the case of
Entomostraca in which these processes are less developed as in
Cyclops or Bosmina.
The numbers of individuals are too small to delineate accurately
the recurrent pulses which are suggested in the data of distribution.
In the autumns of 1895 and 1897, when the occurrences are most
continuous, the larger numbers tend to fall at the times of the
maxima of pulses of other Entomostraca. There is no marked
limitation placed upon this species by the seasonal changes in tem-
perature. It is found throughout the seasonal range in tempera-
tures, though numbers are slightly smaller in channel waters in
November-December. Nevertheless it occurs in considerable num-
bers in the backwaters in breeding activity under the ice at mini-
mum temperatures in December.
282
Of the total individuals, 40 per cent, were males; 45 per cent.,
females without eggs; and 15 per cent., females with eggs. The
sexes show no marked or constant seasonal differences in distribu-
tion. Females with eggs are more abundant in August-October,
and with spermatophores in the same months. Detached sperma-
tophores were found until December.
This species is stated by Herrick ('84) to be distributed in the
entire Mississippi Valley. Marsh ('93) finds it in Wisconsin, but
it appears nowhere in the plankton of the Great Lakes. Brewer
('98) reports it in the backwaters of the Platte in Nebraska, and
Schacht ('97) states that it is an exceedingly common species in
central Illinois, and that it has been reported from Wisconsin, Ohio,
and Minnesota. It thus appears to be limited to the shallow and
relatively warm waters of the prairie regions of the Mississippi basin.
Diaptomus siciloides Lilljeborg. Average number, 10; in 1897,
350; in 1896, 56; in 1895, 282; and in 1894, 23. As will be seen
on comparison, these yearly averages are very similar to those of
the preceding species with the exception that the development of
D. siciloides is about twice that of D. pallidus in 1895. In other
particulars its seasonal data so resemble those of D. pallidus as to
make their discussion in large part a repetition. Its seasonal-
distribution relations to temperature and hydrographic conditions,
breeding season, and its tendency toward a pulse-like recurrence in
coincidence with other Entomostraca are all very similar to these
features in D. pallidus. The proportions of the sexes differ slightly,
the males being less numerous (3 1 per cent.) and egg-bearing females
more abundant (18 per cent.) than in the previous species.
This is also an American species, reported thus far only from
Lake Tulare, Calif., the Illinois River, and waters of Indiana and
Iowa (Schacht, '97), and by Brewer ('98) in lakes and pools of
Nebraska. It is thus confined largely to shoal and warm waters.
Diaptomus spp., immature. Average number, 19; in 1897, 560;
in 1896, 158; in 1895, 336; and in 1894, 120.
The immature individuals of D. pallidus and D. siciloides were
not distinguished from each other in the records. Young Diaptomus
presumably belonging to these two species occur in every month
but March, though but 10 of the 74 records were made in January-
June. The percentage of occurrences and the numbers per m. 3 are
lowest in these months, not rising above 33 per cent, and 500 per
283
m. 3 save in two instances. Occurrences of small numbers continue
through July, but from August 1 to October 15 appear the major
pulses of the year, attaining an amplitude of 1,000 to 8,800 per m. 3
With the decline of temperatures in October, numbers fall to levels
below 400 per m. 3 , with one exception (December 14, 1897) at
700. The percentage of occurrences is, however, high (41 to 44
per cent.) and declines only to 33 per cent, in January. The period
of greatest numbers of young thus coincides with that of greatest
abundance of adults, and lies at temperatures of 70Vand above, in
channel waters.
The effect of hydrographic changes upon the occurrence of
young Diaptomus appears in striking form in the annual averages
above quoted. In 1898, a year of sudden changes, the average
per m. 3 is only 19, while in the stable conditions of the previous year
it is 560. The July-December production in 1897 is 28 times
greater than that of 1898. In 1896, a year of recurrent but less
sudden floods, the average (158) is less than that of 1895 (336), a
more stable year. The great reduction of adults noted in 1898 and
1896 is thus paralleled by an even greater reduction of the young.
Osphranticum labronectum Forbes occurs in the plankton of
Quiver Lake in small numbers (see Schacht, '98), and was found
once in channel plankton in June, 1896.
AMPHIPODA.
Allorchestes dentata (Sm.) Faxon. This is an abundant littoral
species found amid vegetation, especially in the vegetation-rich
backwaters, such as Quiver Lake. It was not often found in channel
plankton, being taken only in the summer of 1895, when the July-
August floods carried away the vegetation which had accumulated
during the antecedent low water.
ARACHNIDA.
ACARINA.
In vegetation-rich backwaters members of the family Hydrach-
nid& were frequently taken, along with other adventitious or-
ganisms, with the plankton. In channel waters they are less
frequent, and are represented principally by Atax, which is parasitic
284
in great numbers (see Kelly, '99) in the Unionida which are found
in the bottom of the channel. Occurrences in the plankton were
limited to the months of May-August, and may be due in part,
especially in the warmer months, to the release of the parasites by
the death and flotation of their hosts. Flood waters in warm
months were often disastrous to the Unionidce because of the load
of silt, sewage, and industrial wastes which they carry in channel
confines at the lower river stages often prevailing in these months.
Other small aquatic Acarina were also present, probably adven-
titious from the littoral or bottom ooze. With two exceptions their
occurrences in the plankton were all in warmer months, April-
September, though not in flood waters. During the period of the
migration of waterfowl, parasitic Acarina were noted in plankton
collections in a few instances.
TARDIGRADA.
Macrobiotus macronyx Duj. Average number, 11. This species
is found principally in the colder part of the year, from October to
May. The earliest autumnal record was October 30, 1895, at 45,
and the latest vernal one, May 1, 1896, at 68.8, and the maximum
number (2,980 per m. 3 ) was recorded on April 10, 1&96, at 46.2. Of
this number, one sixth were females with eggs. Females with eggs
were also found in November, February, and March. Because of
its seasonal distribution it is found principally, though not solely, in
disturbed hydrographic conditions, and its occurrence in the plank-
ton is largely adventitious.
HEXAPODA.
Owing to the shoal waters, relatively narrow confines, and the
hydrographic fluctuations in our fluviatile environment, the aquatic
insects, both larval and adult, have many points of contact with
the plankton. They constitute a large element in the total volume
of the animal population of shore and bottom, and are all connected
by chains of food relations, more or less complex and remote, to the
plankton organisms or their sources of food. With the single
exception of the larvae of Corethra they are all in the main adventi-
tious members of the plankton assemblage, and are much more
abundant in the vegetation-rich backwaters than in the channel.
285
Since the aquatic insects of these collections are being studied by
others, with reference to publication in this Bulletin (see Hart, '95,
and Needham and Hart, '01), only passing notice of the more
important representatives appears in this connection.
EPHEMERIDA.
Ephemerid larvae, as a rule in early stages, were found singly
or in small numbers in the channel plankton in the warmer months,
April-October, at temperatures above 56. Since these occurrences
were with few exceptions in stable hydrographic conditions, it seems
probable that the younger larvae of this order may adopt, at least
temporarily, a limnetic habit. Specific identifications of these
larvae were not made.
HEMIPTERA.
Corisa (?) sp. Average number, 37. A small hemipterous
larva doubtfully referred by Mr. C. A. Hart to Corisa, was taken
with some frequency but in relatively small numbers in the plankton
during the summer months. Of the 36 occurrences 27 fall in
June-August, 2 in May and 3 in September, 2 in January, and 1 each
in October and November. It thus appears in the temperature
extremes, but exhibits a great predominance in the season of maxi-
mum heat. There is no marked increase in its frequency or numbers
in years of more disturbed hydrographic conditions. Its numbers
are always small and somewhat erratic. Adult Corisa, as well as
many other aquatic Hemiptera, were found in plankton collections
singly and infrequently.
DIPTERA.
This group of insects is abundantly represented in the plankton,
but in all cases by larval or pupal stages.
Chironomus spp., larval stages. Average number, 124. Larvae
in various stages of development from that immediately after
hatching to that approaching pupation were found in channel
plankton. They occur in considerable numbers in the ooze in the
river bottom, but appear to abandon the limicolous for the limnetic
habit, temporarily at least, as a result of hydrographic or other
disturbances. There is evidence from their relative numbers in
286
years of different hydrographic conditions that these have consider-
able influence in bringing them into the plankton. Thus in 1897,
in stable conditions, there were only 5 occurrences in 3 1 collections
examined, averaging 88 per m. 3 , while in 1898, in more disturbed
conditions, there were 29 occurrences in 52 collections, averaging
124 per m. 3 There is also a marked seasonal distribution. The
larvae appear in the plankton in March-December through the
seasonal extremes of temperature, but the numbers in March and
November-December are always small. Only 15 per cent, of the
occurrences and 5 per cent, of the individuals were found at tem-
peratures below 45. The percentage of occurrences in the collec-
tions is highest in March-September, the percentages being 53, 73,
80, 47, 78, 52, and 50, respectively, to 8 to 35 per cent, during the
remaining months.
Corethra sp., larval stages. Average number, 6. These semi-
transparent and active larvae have the characteristics of limnetic
organisms, and may be reckoned among the autolimnetic planktonts
of our waters. Because of their activity , it seems probable that
they escape the drawn net, especially the small model used by
us, and also, because of their negative rheotaxis, elude the suction
of the plankton pump to an even greater extent. Thus, in 1895, in
net collections, there were 8 occurrences averaging 32 per m. 3 to 4
in 1898, in pump collections, averaging 8 per m. 3 Corethra larvas
were never abundant in our plankton, probably in part for the
reasons just cited. With two exceptions all the occurrences lie in
the period of maximum temperatures in June-September, 7 of the
14 occurrences and one third of the individuals being recorded in
August.
Dixa sp., larval stages. Average number, 8. Larvas were
recorded singly in scattered occurrences in all months but February
and October-December, though most of them appear during maxi-
mum temperatures.
Larvae of Tanypus and Odontomyia were also recorded in May
and June in isolated occurrences.
In addition to the larval stages of these aquatic insects there
occurred in the plankton a considerable number of insect eggs,
principally those of Diptera and Ephemerida. These were generally
isolated, though sometimes fragments of the egg-string of Chirono-
287
mus appeared. They were recorded in all months but February
and December, though 20 of the 30 records and 81 per cent, of the
individuals appeared in May- August. The numbers are never very
large, the maximum record, 5,424 per m. 3 on June 29, 1894, being
due to a number of fragments of egg-strings.
MOLLUSCA.
GASTROPODA.
The adults and young of many of our aquatic gastropods have
the habit of gliding on the under side of the surface film of water, and
they are also frequently dislodged from their foothold on aquatic
vegetation, and thus enter the habitat of the plankton temporarily.
This is especially true in vegetation-rich backwaters. The smaller
forms, such as Ancylus, Amnicola, and Planorbis parvus were occa-
sionally taken in the summer plankton of the channel.
LAMELLIBRANCHIATA.
This group is represented in the plankton by the larval stages,
or glochidia, of the Uniomdcz, which form an important part of the
bottom fauna of the stream and its tributaries.
Anodonta corpulenta Cooper. Average number of glochidia, 21.
The seasonal distribution of the glochidia in the plankton is very
well defined. With but two exceptions the 48 occurrences all fall in
October- April, and 40 of them in November-March. The occur-
rences are thus during the period of minimum temperatures ; indeed,
31 of the 48 are at temperatures not exceeding 35 in surface waters,
and only 9 are above 45. The earliest autumnal record is Septem-
ber 30, at 58, and the latest vernal one, June 6, at 79. Generally
the earliest records are in the closing days of September or the early
ones of October, and the latest records are about the first of April.
The occurrences are more frequent in December-March, the glo-
chidia appearing in 64, 50, 53, and 60 per cent, of the collections,
respectively, in these months. Their numbers are also several
fold greater at this season than in the earlier and later months of
their occurrence. The period of minimum temperatures is thus
the season of greatest discharge of glochidia. The numbers are
always relatively small, 520 on December 28, 1897, being the maxi-
288
mum record. Their fluctuations are erratic, and show no apparent
relation to hydrographic or other environmental changes.
Lampsilus anodontoides (Lea) Baker. Glochidia referred with
some uncertainty to this species appeared somewhat irregularly in
the plankton in small numbers in September-December and again
in June-July. The seasonal distribution in two periods suggests
the inclusion of two species.
Arcidens confragosus (Say) Simpson. Glochidia of the type
referred by Lea to the old genus Margaritana, and presumably
belonging to this the commonest member of this genus (as formerly
understood) in our locality, were taken in the plankton December
18, 1895, in small numbers.
BRYOZOA.
This group is represented in our plankton by the floating stato-
blasts, when these occur, as in Pectinatella and Plumatella, by
detached and floating fragments, as in Urnatella, or by natant
colonies, as in Lophopus and Cristatella. Genera such as Fredericella
and Paludicella, whose statoblasts sink, fail to appear in the plank-
ton, though in some cases they may be abundant in the bottom
fauna. The Bryozoa are plankton feeders, and play an important
role as plankton reducers in vegetation-rich backwaters.
DISCUSSION OF SPECIES OF BRYOZOA.
Cristatella mucedo Cuvier. This species was found in the back-
waters in summer months, especially in Quiver Lake. Statoblasts
probably referable to this species occurred sparingly in May and
August.
Lophopus cristallinus Pallas. This rare bryozoan occurred in
the channel plankton, though not in our quantitative collections, in
July, 1897, in that part of the channel containing the discharge from
Quiver Lake. Small, free-swimming colonies of 5-50 zooids were
taken in surface waters.
Pectinatella magnified Leidy. Statoblasts of this superb bryo-
zoan were not uncommon in the backwaters, and were seen several
times in the vernal plankton of the channel. The large floating
colonies are found near the surface in July-October in the open
backwaters, and more rarely in the river itself. The translucent
289
gelatinous coenoecia are spherical, ellipsoidal, or often somewhat
flattened. The longest diameter of these floating masses often
exceeds 30 cm.
Plumatella repens L. This is by far the most abundant bryozoan
in our locality, being found everywhere on submerged vegetation
in the backwaters. It often develops with surprising rapidity on
the submerged stems of plants, where, as in 1896, summer floods
reinvade the vegetation-covered margins of reservoir backw r aters.
It is represented in the plankton by its floating statoblasts. Their
seasonal distribution shows some correlations with temperature,
hydrographic conditions, and the seasonal cycle of the parent
organisms. During the period of minimum temperatures (Decem-
ber-February, inclusive) they are relatively rare in the plankton,
appearing in 30, 8, and 20 per cent., respectively, of the plankton
catches. They are rare in high- as well as low-water conditions, as,
for example, in the floods of 1895-96 and 1898, when they appear
in but one of 15 collections. With the rise of temperature in March
they occur more frequently, as, for instance, in 1898 (Table I.), and
continue during the run-off of the spring flood. The occurrences
rise in March-May to 60, 46, and 50 per cent, of our total collections
in these months, and the numbers also are larger. For example, in
1898, 81 per cent, of the total individuals for the year were found
in these months. The discharge from impounding backwaters, the
principal breeding grounds of the parent organisms, doubtless tends
to increase the numbers of statoblasts in channel plankton during
this season. During the remainder of the year, June-November,
the percentage of occurrences again falls to 30, 50, 24, 32, 18, and
44 per cent., respectively. The 50 per cent, in July is due to the
summer flood of 1896. If this year is omitted the record falls to
33 per cent. The large percentage for November is probably due to
the predominantly higher levels of this month, to the invasion of
lake margins seeded with statoblasts, and to the increased activity
in the fishing industry, which tends to disturb the summer's growth
of vegetation in tributary backwaters. The relations to the seasonal
cycle of the species are patent. The summer months, June-
September, are the season of growth and spread of the parent
organisms and of the formation of statoblasts, especially as receding
levels expose the water margins. Hydrographic or other disturb-
ances tend to increase the number of statoblasts in the plankton
290
until minimum temperatures are reached, when minimum numbers
appear in the plankton. As temperatures rise, the statoblasts
tend to float and become more abundant in the plankton, as a result,
perhaps, of the physiological and accompanying physical changes
in the contents of the statoblast. The declining phase of the major
flood of the year is thus the period of greatest flotation and dispersal
of the statoblasts.
Urnatella graciUs Leidy. This unique species is found in some
abundance on the projecting margins of the shells of the Unionida
which line the river bottom in many reaches of the channel. Small
fragments of the colonies containing only several polypides were
found in the plankton in May-August and October. The earliest
record was May 25, and the latest, October 25, at 48.5.
THE PERIODICITY IN THE MULTIPLICATION OF THE ORGANISMS OF
THE PLANKTON.
One of the most obvious conclusions brought to light by the
detailed study of the volumetric fluctuations of the plankton pub-
lished in Part I. of this report, and most strongly reinforced by the
statistical data showing the fluctuations in the numbers~of the indi-
viduals of the various species and in the sums total of the various
biological groups represented in the limnetic fauna and flora, is that
plankton production is fundamentally rhythmic or periodic in
character, viewed either in its constituent elements or as a whole.
This total result is simply the sum of a like phenomenon pervading
more or less completely and coincidently the reproductive cycles,
the rise and decline in the numbers of the typical constituents of
the plankton. The exceptions to this rhythm are usually found in
those organisms which are adventitious in the plankton and have
their centers of growth and distribution in other regions than the
open water.
Many illustrations of this periodic movement in the multiplica-
tion of organisms of the plankton have been cited in the preceding
pages and may be seen in the accompanying plates. As an illustra-
tion for discussion in detail we may take the pulse of July, 1898,
shown in the volumetric data of Table III. and Plate XII. of Part I.
The fluctuations in the biological population during this period are
also tabulated in Table I. of this paper, and graphically presented
in Plates II. and IV., which exhibit the movement in the totals of
the Chlorophycecz, Bacillariacece, and chlorophyll-bearing Masti-
gophora, and of the Rotifera and Crustacea.
In the volumetric data the pulse rises from a minimum of .14 cm. 3
per m. 3 on July 5 to a maximum of ,88 cm. 3 on the 19th, declining
again on the 26th to the second minimum, of .67 cm. 3 Its duration
is thus four weeks and its amplitude, in comparison with many
other pulses in the records, relatively slight. It occurs in the more
stable conditions of declining river levels and midsummer tempera-
tures. The following list gives the names of the more or less typical
planktonts considered in the discussion of this pulse. Others,
largely adventitious or insignificant in numbers, might be added
(20) 291
292
to the list. Forms whose antecedent minimum does not fall on
June 28 or July 5 are designated by a superior 1 ; those whose
maximum does not fall on July 19 or 26, by a superior 2 ; and those
whose subsequent minimum is not on July 26 or August 2, by a
superior 3.'
The component forms and groups are Crenoihrix, etc. 1 , total
Schizophyce, Microcystis ichthyoblabe 1 , total Chlorophycea, Actinas-
trum hantzschii, Crucigenia rectangularis, Pediastrum boryanum 1 ' 2 ' 3 ,
P. pertusum 2 ' 3 , Raphidium polymorphum 1 , Scenedesmus genuinus,
S. obliquus, S. quadricauda, Schroederia setigera, total Bacillariacece 1 ,
Cyclotella kuetzingiana, Diatoma elongatum 1 , Fragilaria virescens 2 ,
Melosira granulata var. spinosa 1 , M. varians 2 , Navicula spp., Synedra
acus, total Conjugates 1 , Closterium acerosum, C. gracilis, total Protozoa,
total Mastigophora, Eudorina elegans, Euglena acus, E. oxyuris,
E. viridis, Glenodinium cinctum, Lepocindis ovum, Pandorina
morum 3 , Phacus longicauda 1 - 2> 3 , P. pleuronectes 2 ' 3 , Platydorina
caudata, Pleodorina calif ornica, Trachelomonas acuminata 1 , T.
hispida 3 , T. volvocina, total Rhizopoda, Difflugia globulosa, total
Ciliata 2 , Codonella crater a 2 , Halteria grandinella 2 * 3 , Tintinnidium
fluviatile 2 ' 3 , total Rotifera, total Bdelloida 1 ' 2 , total Ploima, Anurcsa
cochlearis and var. tecta, eggs of A. cochlearis and var. tecta, A.
hypelasma, Asplanchna brightwellii 1 ' 2> 3 , Brachionus angularis and
var. bidens, eggs of B. angularis and var. bidens, B. bakeri and vars.
duniorbicularis 1 , melhemi, and tuber culus 1 ' 2 , total of all varieties
of B. bakeri, B. budapestinensis, B. militaris 1 ' 2 , B. pala and var.
amphiceros, B. urceolaris var. bursarius, B. variabilis 2 * 3 , Mastigocerca
carinata 1 , Monostyla bulla, Polyarthra platyptera, eggs of P. platyp-
tera 2 ' 3 , Rattulustigris 2 , Synch 3 , Bosmina longirostris 1 ' 3 , Ceriodaphnia scitula,
Chydorus sphcericus,Diaphanosoma brachyurum 3 ,Moina micrura 1 ' 2 ' 3 ,
total Copepoda lj 2> 3 , Cyclops viridis var. brevispinosus and var.
insectus, C. edax, young Cyclops 1 , nauplii of Copepoda 1 ' 3 .
An examination of the preceding list and of the qualitative data
of Table I., reveals the fact that 71 of the more typical planktonts
are found in appreciable numbers in the plankton during this
month. To this number we may add 6 immature forms separately
listed in the table and 14 group totals, making in all 91 sets of
statistical data bearing on the components of this pulse. An analy-
293
sis of the behavior of the constituent species shows that 43 of the 7 1
species (including varieties and forms), 4 of the 6 immature forms,
and 10 of the 14 group totals reach their greatest amplitude on the
19th, coincidently with the volumetric maximum. Thus, in all, a
total of 57 out of 91, or 63 per cent., of the sets of data are in pre-
cise agreement as to the time of maximum development. Fur-
thermore, of the remaining 35, there are 10 culminating in the
collection prior to the 19th (on the 12th), and 16 on the next subse-
quent one (on the 26th,) in all, 26 or 29 per cent, which culminate
on immediately contiguous dates of examination. This leaves a
residuum of only about 8 per cent, which do not exhibit precise or
substantial agreement as to the time of maximum development. In
the matter of the location of antecedent and subsequent minima the
agreement is less pronounced, possibly because the enumeration
error is relatively greater in the case of minimum numbers. We
find, however, that 65, or 72 per cent., of the antecedent minima
of the pulses occur on June 28 or July 5, and 71, or 79 per cent., of
the subsequent minima are on July 26 or August 2. Nineteen, or
20 per cent., of the antecedent minima are on July 12; and 10, or
11 per cent., of the subsequent ones are on August 12. There is
thus a residuum of not over 10 per cent, of instances where the data
of species or group totals do not coincide or approximate to this
pulse, as described, in position of maximum or one or both of the
limiting minima. Considering the necessarily large error entering
into our data, it is not surprising that exceptions should occur.
Some exceptions as, for example, that of Pediastrum pertusum
(Table I.) are plainly not due to insufficient data, but are appar-
ently normal dislocations; that is, the rhythm of this species at this
time is not in harmony with that of the majority of the components
of the plankton. . But this is only a temporary derangement, and
is not the habitual relationship which movement of production in
Pediastrum bears to that of the plankton as a whole. So, also,
many of the Entomostraca are much delayed in the culmination of
their increase, running over to August 2 or 9, while the most of the
other planktonts culminate on July 19 or 26. This lag on the
part of the Entomostraca is not, however, habitual, as will be seen
on examination of Plates II. and IV. This tendency toward a
coincident rhythmic movement in production on the part of the
constituent organisms of the plankton will be found throughout all
294
the data where collections are of sufficient frequency to adequately
delineate the curve of production, that is from July, 1895, to Oc-
tober, 1896, and from July, 1897, to March, 1899, a total of 37
months, and suggestions of a like phenomenon appear in the less
complete data of other years. The degree of agreement indicated
in the pulse of July, 1898, will be found, on examination of the data
in Table I. and in the plates of this paper, to vary with the environ-
mental conditions. Times of rapid change in hydrographic condi-
tions or in temperature generally show less agreement, and more
stable conditions will exhibit an equal or even greater uniformity
in the prevalence of the pulse-like rise and decline of the component
organisms.
In order to show the course of these recurrent pulses in the
chlorophyll-bearing planktonts, the total Chlorophycetz, Bacil-
lariacecE, and chlorophyll-bearing Mastigophora on the one hand, and
of the Rotifera and Entomostraca (''Crustacea" of the plates), I have
presented the data graphically on Plates I.-IV., and in the table on
pages 296-299 have drawn up a list of the pulses, indicating the
dates of the collections which in the main enter into the respective
pulses, and the dates of the maxima or culminations of the five
groups named. Owing to the irregularities in the data, there are
some instances in which several possible dates might have been
chosen. Reasons for the choice are in several important instances
given in the foot-notes 'to the table.
It is evident from the data here presented in graphic and tabular
form that the pulses of the five groups of organisms tend in the main
to coincide. This is shown in Plates I.-IV., and in the fact that the
average divergence of 175 group pulses listed in the table is 6.4
days, or, if 5 aberrant instances are omitted, only 4.8 days. In other
words, the pulses of the totals of the 5 groups included in the table
culminate on an average within an interval of 6.4 (4.8 in 170 cases)
days. The average of the extreme limits between maxima of
group pulses in the 36 periods of movement listed in the table is
11.7 days.
It is apparent that the pulses would be more completely de-
lineated by collections at daily intervals, but even in the somewhat
irregular and at times chaotic data here presented, the evidence
seems conclusive that the seasonal production of the dominant
species and groups of planktonts tends to fall into coincident
295
recurrent pulses, which, in turn, are the cause of the similar and
often coincident volumetric fluctuations.
Attention should be directed to the fact that without any im-
portant exceptions this recurrent movement pervades all the
organisms of the plankton which are eulimnetic, such as Scenedes-
mus, Melosira, Trachelomonas, Codonella, Synchtzta, Daphnia, and
Cyclops, and often those which at certain seasons become tempo-
rary planktonts, such as Difflugia and Hydra, but not with any
regularity the tycholimnetic organisms, such as bdeltoid. rotifers or
nematodes. It affects the more highly organized Rotifer a and
Entomostraca with slower growth, longer life, and consequent
greater cumulative function as well as the algae, diatoms, and
flagellates, where rapid multiplication, brief existence, and non-
cumulative (in the individual) function prevail. The large share
which the young (eggs and immature stages) play in the pulses of
Rotifera and Entomostraca will be seen in Table I., and repeated
attention has been called to this in the discussion of species. The
prevalence of breeding females and of eggs or young during the rise
of the pulse, and of eggless, moribund, or dead individuals or their
skeletons during the decline, is a common phenomenon in all well-
defined pulses. No species of plankton organisms appears to escape
the operation of this recurrent movement in production.
The proportion of individuals surviving from one pulse to the
next is subject to great variation, being often least when the ampli-
tude of the pulses is greatest, and largest when the pulses culminate
at slight amplitudes. As a result of periods of minimum develop-
ment, it follows that the possible length of life of most plankton
organisms, even of the Rotifera and Entomostraca, in the plankton
must fall within rather narrow limits of a few days or a fortnight
at the most. Since the contrasts between minimum and maximum
numbers are relatively greater among the chlorophyll-bearing
organisms, it follows that the survival proportion is less in these
groups.
The duration and amplitude of the plankton pulses will vary
within certain limits according to the method of delineation. The
volumetric minima and maxima present the total product in cubic
centimeters, and the pulses thus marked cat have been described
in Part I. They may also be delineated by statistical data of the
total plankton or of its larger groups of organisms, or by the domi-
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300
nant or more typical species. In the case of the total plankton
some obscurity results at times from the inclusion of unusual
proportions of -an adventitious population with flood waters. The
selection of particular organisms as representative is also subject to
some error, since seasonal changes in temperature and other more
subtile causes often deflect or suppress their development. The totals
of the ChlorophycecB, Bacillariacece, and chlorophyll-bearing Masti-
gophora, and of the Rotifera and Entomostraca (PL I. -IV.) probably
give as complete and accurate a delineation of the recurrent pulses
as the statistical data afford, since they include relatively few
adventitious organisms, cover the entire year, and swamp more or
less completely individual and temporary divergences of particular
species. The delineation of the pulses by statistical data is obviously
more significant than the volumetric method, since it more clearly
presents the results of the reproductive processes which lie at the
foundation of the phenomenon of recurrent pulses ; and this method
is also free from the unavoidable error arising from the presence of
silt in the collections.
The interval between collections introduces an error of consid-
erable moment in any effort to determine with accuracy the duration
of individual pulses, that is, the length of time between their minima
or maxima. Daily collections would render this feasible, but with
an interval of a week or more, not only the duration, but in some
cases the probable separation of the pulses and location of their
maxima, is to some undetermined degree obscured.
The duration of the pulses of the five groups of plankton organ-
isms shown graphically on Plates I. -IV., in the case of all chlorophyll-
bearing organisms considered as a whole, is in 29 out of 36 instances
between 21 and 35 days, less than 21 in 2 cases, and more than 35
in 5, reaching extreme limits of 14 and 49 days. They average
30.25 days between minima and 29.97 between maxima.
The rotiferan data in the same months may be divided into 36
periods, in 33 of which pulses are traceable. The duration of pulses
between minima lies between 2 1 and 3 5 d ay s in 2 3 of the 3 6 instances ,
falls below 21 in 5, and is above 35 in 8. The extreme limits are
14 and 49 days.
In the case of the Entomostraca, where also the pulses are obscure
in a few of the intervals, we find that 22 of the 36 are between 21
and 35 days between minima, 5 are below 21, and 9 are above 35.
301
The extreme limits are 12 and 49 days, and the average duration
is 29.9 days.
From the data here presented it is evident that the pulses are in
the main from 3 to 5 weeks in duration, averaging approximately
29 + days a little less than one calendar month.
The amplitude of the pulses is affected profoundly by seasonal
and local influences, such as the factors of temperature and chemical
constituents of the water, and the hydrographic conditions. These
have been discussed in connection with the volumetric-data in Part I.
and in the discussion of species in the first part of the present paper.
Rising, or even uniform, temperatures, hydrographic stability,
decaying vegetation or access of sewage or other fertilizing constitu-
ents, all serve to increase the amplitude of the pulses. Declining
temperatures, dilution or suspension of access of fertilizers, competi-
tion of gross vegetation, access of flood waters and increase in
current, all tend, in the main, to depress the amplitude of the pulses.
The duration of the pulses is not, however, thereby essentially
modified, though a tendency to override subsequent pulses and
partially, rarely wholly, to submerge them is at times of major pulses
often apparent in the data.
The cause and significance of the phenomenon of recurrent pulses
is not clearly and unmistakably evident, owing, on the one hand, to
the irregularity of the data, and, on the other, to the great complex-
ity of the problem, especially in the fluctuations and varying
combinations of environmental factors.
The plankton method itself is subject to great errors, but these
are largely distributed, and careful examination, especially of the
matter of dilution and computation, has failed to reveal any probable
or even possible source in the method to which these recurrent pulses
can be traced.
It is not impossible that the rhythm here noted is merely a
chance outcome of the statistical method and without biological
significance; that it is wholly accidental, the resultant of the con-
flicting and varying factors of the environment and not predomi-
nantly or continuously initiated by any one factor. On the other
hand, its nature, as we have described it, is such that we are led to
look for some factor in the environment with which this rhythm of
repetition in growth of the plankton organism might be correlated, or
to some internal or inherent factor within the organisms constituting
302
the plankton, or to the interaction of environmental and internal
factors.
That there is a periodicity in the reproductive processes of
organisms, of both plants and animals, is generally apparent. We
see it in the flowering and fruiting seasons of the phanerogams, and
in the breeding seasons of many invertebrates, of mollusks and
insects, and of the vertebrates generally, of fishes, amphibians,
reptiles, birds, and most mammals. Fluctuations in environmental
conditions, notably in food and temperature, influence these re-
productive processes. The phenomenon of rise and decline of the
microscopic population in laboratory aquaria is likewise an illustra-
tion of the periodicity of organisms, but usually within a briefer
interval than that of the organisms above mentioned. The studies
of Maupas ('88) and Calkins ('02) have shown that even in the
seemingly uniform conditions of the laboratory, the reproduction
of the ciliate Protozoa is essentially periodic.
On a priori grounds it seems highly improbable that in the case
of the organisms of the plankton, internal factors should determine
the coincidence of the periods of growth and reproduction in several
hundred species. While it is not impossible, or indeed improbable,
that these species of the plankton if bred in pure cultures or uniform
environment would still exhibit a periodic reproduction, it seems
highly improbable that so diverse an assemblage of algae, diatoms,
flagellates, protozoans, rotifers, and entomostracans as is found in
the Illinois River, would exhibit in laboratory cultures under
uniform conditions any such coincidence in the location and duration
of their pulses as is found in the waters of the stream. Whatever
the internal factors involved in the growth and reproduction of
plankton organisms may be, it is patent that we must look for some
environmental factor or factors lying at the foundation of the
coincidence of seasons of growth and reproduction of plankton
organisms, which results in the phenomenon of recurrent pulses in
species, groups, and volumetric plankton.
We may simplify the problem somewhat by recognizing at the
outset the importance of nutrition in supplying the basis for the
periodic growth of any organism. The rotifers and entomostracans,
at least the limnetic types, depend in large measure, either directly
or indirectly, upon the synthetic planktonts, such as the algae, dia-
toms, and flagellates, for their food. Since the pulses of these animal
303
forms (cf. Plates III. and IV. with I. and II.) coincide with or
follow shortly after those of the synthetic planktonts on which they
feed, we may conclude that the cause of the periodic movement of
these animal groups lies in the periodic fluctuations of their food
supply. In the causes which control this periodic growth of the
chlorophyll-bearing organisms will be found the solution of the
general periodic phenomenon in plankton.
This rhythm is primarily one of growth and reproduction, and
its solution must be sought in the forms of matter anctenergy which
affect these processes. The nutrition of the chlorophyll-bearing
organisms is drawn from matter in the river water. The analyses
contained in Part I., Table X., and graphically presented on Plates
XLIII. to XLV. trace the seasonal fluctuations in the nitrates -one
of the important constituents of plant food. Neither in the seasonal
curves of this or other forms of nitrogen delineated in the plates is
there any such rhythm of occurrence, though, as has been pointed
out in the discussion of the chemical conditions, there are instances
of apparent correlation of plankton and nitrate pulses. They occur
at irregular intervals, and do not form a continuous series. That
there might be a rhythm in the utilized nitrates (the analysis repre-
sents only the unused residuum) is of course possible, or that it
might occur in some other constituent of the food not determined
in the analysis is not impossible, but we have no evidence of its
existence.
The chlorine in our river waters is a fair index of the amount of
sewage or pollution by animal wastes. It is subject to considerable
fluctuations, resulting in part from dilution by floods or concentra-
tion in low waters, and there are other pulses not traceable to
hydrographic conditions, which perhaps result from industrial
wastes. These fluctuations in some instances coincide with those
of the phytoplankton in question, but the instances are few and
the correlation is incomplete. Upon investigation I find that
sewage pumpage at Bridgeport, which discharged the sewage of
Chicago River into the Illinois and Michigan Canal and thence into
the Illinois River, was practically continuous, and could not produce
the rhythm 'in question. The sewage of Peoria has a much more
immediate effect upon the chemical conditions in the river at
Havana than has that of Chicago. The sewers of this city, I am
informed by Mr. H. E. Beasley, City Engineer, are flushed as
304
follows : " The method used is that of flushing with a hose, a crew
of men being kept constantly at work, taking them about a period
of three weeks to cover the entire system. The water is allowed to
run through a fire-hose at each point for a period of about ten
minutes." This system was in use during the years of our opera-
tions, and it offers no occasion for the periodic pulses in growth of
the organisms in question. Investigation of the discharges of dis-
tillery and cattle-yard wastes into the stream has not revealed any
periodic fertilization of the river waters from these sources. The
available data thus fail to exhibit any periodic rhythm in food
matters in solution and suspension in the river water with which
these pulses of chlorophyll-bearing organisms might be correlated.
Frequent reference has been made in previous pages to the
appearance of pulses upon the decline of floods. Flood waters bring
into the river, as shown by the chemical analyses, large quantities of
silt and organic wastes in suspension and solution. They inundate
great tracts of fertile territory rich in vegetation, and thus add to
the available sources of food for the phytoplankton. Decline of the
flood affords time for decay and solution of some of the food matters,
and time also for breeding, and its run-off adds to the volume of the
plankton in channel waters. A comparison of the hydrographs of
the years in question (Part I., PL X.-XIII.) with these recurrent
pulses (PL I.) will show that many if not most of the pulses appear
on declining flood waters, and that many of the larger ones follow
the major floods. Closer analysis, however, shows that there are
sometimes two pulses of chlorophyll-bearing organisms on the
decline of a single flood, and that they may also occur upon rising
flood or even in its entire absence. Floods unquestionably affect
the amplitude of the pulses, and to some extent modify their location.
They seem inadequate, however, to explain their recurrence and
their tendency toward a uniform interval. Minima between pulses
also recur on declining floods.
Energy as well as matter is necessary for the growth of the
phytoplankton, and its source is primarily the radiant energy of
the sun. A plot of the tri-daily air temperatures at Havana for
1894-1896 (Part I., p. 478, Fig. C) inclusive, exhibits many irregu-
larities, a few of which partake of the nature of recurrent pulses at
approximately monthly intervals, but they are too few and too irregu-
lar to be the basis of the recurrent growth of the phytoplankton.
305
The importance of light for the photosynthesis of chlorophyll -
bearing plants is unquestioned. The liberation of oxygen by the
plant declines as the light fades, and is at its lowest ebb in darkness.
The access of light to the phy toplankton is limited by several factors
of the environment, principally by silt, which increases the turbidity,
and by clouds, which interfere with the penetration of the sun 's rays.
The fluctuations of the silt are chiefly the result of floods, and, as
above stated, the floods do not exhibit a rhythmic pulse which can
be correlated with that of the phytoplankton ; much less do the
periods of rising water which are most silt-laden. The cloudiness
of the sky varies greatly at different seasons of ' the year, being
predominant at times in the autumn or winter months. It is sub-
ject to pulse-like occurrences of variable duration, but an examina-
tion of the records for central Illinois for the years under discussion
does not disclose any periodic rhythm which can be correlated
continuously with that revealed in the statistical records of the
growth of the phytoplankton.
Another factor of the environment which modifies the quantity
of light which impinges upon the chlorophyll-bearing organisms
of the plankton is the light from the moon. The amount of light,
both absolute and relative, derived from this source is not great.
According to the calculations of Zollner, the light from the sun is
618,000 times as bright as that from the full moon. In the pre-
sent connection it is only important to know whether the moon-
light contains an amount of solar energy sufficient to appreciably
affect the photosynthesis of the phytoplankton. The amount of
such energy utilized in photosynthesis is relatively a small propor-
tion of the total, so that there is a possibility that moonlight may
contribute to the process to an appreciable extent.
This matter was investigated by Knauthe ( '98), who determined
the fluctuations in the gaseous contents of the w*aters of carp ponds
rich in Euglena. While this author does not report upon the plank-
ton of the ponds investigated, it seems quite probable that carp
ponds rich in Euglena would present conditions very similar to those
found in the Illinois River, which has a remarkably well-developed
Euglena water-bloom, and abounds also in carp.
The following table presents the results of his work bearing
upon the point in discussion.
55
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l^- r-- CN CN
o3 ^"^
O- ON CN CN '-;
t^.
. . . . 03
cj
O O O O r^
"ri O
03
O O ^~ ' ^"^ CN CN ^O 1O f^- t^* vO NO ^ ^ fO CN CO PO ^O l^"
loiom loinio
^
CNCN t^l-^ OO OO CNCN * Tf CNCNCN 't l ^-^t l CNCN
* * * ^H H ^H
J
OO OO -rt r-i CNCN OO OO OOO OOO OC
OOO -^M t i ^-i
0)
9 S a 8 S
^j --2 - Z| o3 O O
fa
o
,d
t/3 ^3 T^H *^ O O
o o -^
"rt
0)
w 2 - ' fl S-I
o - ^ rZ-i ^ C3 x* ^ "S 'K ^
* S K* b/)' "~*
M
?H
S 3 -2 ^ S^^^ ".^T3
Pl,Q
f- 3 4-^ '^ *^~ l 4-> '-' rl "^ rt
-*- 1 O
OTCfl<^
03 o3
O
s s ^
^ ^
3
2 "3 o3
IH U
o
UJ CO
da
rj ;j
0) CO 0) CO CO CO
eS 03
fl El T)
o3 o3 o3 o3 o3 o3
IH t-( In TH t-H IH D u O3
C PI
c5 CTj
33:3 333 o3 03 ft
WC/2C/2 C/3WC/2 C/3 t/2 C/D
CO CO
00
^ s : s : 6 ! r A d e 6 !
^
00
O3o3o3ftft ftp, ft
ro
rorOO OO1J O O O
O ro fO
ft
o
o o
o
CNl/IONlOON'-H - 1
... CN-ON ^H
T-H TjH
^ x >< x x x x"' x" x
KJ kj
rS rN
1 1 I 1 1 1 II 1
^< 10 i/"; 1O ^ ^ CO fO fO
1 1
306
307
The amount of oxygen present in the water in the dark, or on
dark nights, is reported as 0.20, 0.25, and 0.27 cm. 3 per 100 cm. 3 of
water. In bright sunlight in the laboratory, and with the unusual
abundance of Euglena due to the collection of the water sample
from the region of the water-bloom, it rises to 2.05 cm. 3 In the
case of the Spandauer samples it rises from 0.25 in the dark to 1.15
(an increase of 0.90 cm. 3 ) after "long" exposure to bright sunlight
in the laboratory. The oxygen in this water at 11 :00 p. m., after
exposure to moonlight, amounted to 0.45, or 0.20 cm. 3 more than
was found in control water kept in the dark. In this instance the
apparent increase due to moonlight is 2 / 9 of that due to sunlight. In
the case of the moonlight the analysis was made at 11:00 p. m.,
after not more than three hours ' exposure. The moon was not at its
greatest efficiency, since full moon occurred four days prior to the
date of analysis. In the case of the sample exposed to the sunlight
the analysis was made at 4:00 p. m., after "langer intens Sonnen-
schein." It would seem probable that the effectiveness of moon-
light in comparison with sunlight in photosynthesis by the phyto-
plankton here indicated (2 to 9) is below the possible maximum and
also above that of the average, since it was obtained when the
moon was but four days past its maximum effectiveness.
If we accept Knauthe 's data as sufficient to establish the effec-
tiveness of moonlight in increasing photosynthesis, and thus the
growth of the phytoplankton, we find in it a recurrent factor of the
environment to whose influence we may seek to attribute the rhythm
of growth of the chlorophyll-bearing organisms.
On Plates I. and II. I have plotted the seasonal distribution of
the totals of the Chlorophycece, of the Bacillariacecz, and of the
Mastigophora from July, 1897, to April, 1899, and have indicated
the times of full moon throughout this period by marks at the bot-
tom of the diagram. The diagram shows clearly the occurrence of
these recurrent pulses, their approximation in the three groups of
chlorophyll-bearing organisms upon the same or adjacent dates, and
the occurrence of their maxima in some cases at the time of full
moon or within an interval of ten days thereafter.
In the table which follows, I have given the data bearing on the
pulses of the total of all chlorophyll-bearing organisms from July,
1895, to October, 1896, and from July, 1897, to March, 1899, inclu-
sive, 36 months in all, stating the location of the pulse as determined
(21)
*8
\O ** co *"* '"' ***
OOOON^NC 1 '" 1 -*^
r^ CN CN CN 04 ^ CN
I s
03
IM
1" t t ,5 J 3
H^-, < co O *& P Q
r d I* t- *** ^ ^> b
2 fli aj CO l2 ^ *"
rt., C H 3 ^j;
i ,pH^i
Cca +J
W3 (H .^
O c/3 Q) p>
Qs t^* IO QQ ^ to
l-HTfl-rHvOCO'-INOCO
lONQCN-'-lt^OCOOO
' ^-H
^H CN >-c >-! CN --I
S3 8 &
J2 bC
+ + + + + + +
+ + + + + + + +
' ^
3
N
^
>. X
>%
oj *3
Oj 3 3 3 3 ^ O3
CB3333335
Q
^ 'O
a |
O) f~ ON OO "* O "^
LO t^ NO ^^ NO CO IO Tf*
^H CN CN -i CN '-H CN
8 03
1 1
d
,53
5 c
Q
O O 1 O ro -* O O
I^-COIOCNIOCN'-'CN
'So
i i 7 i i i
1 1 1 + + + 1 1
03
"o
S
2 ^ 2 - ,0 - o
^-cs2-2cx>-
1
^> bii "^ .j ^ d d
"33 o O 0) OJ
^ < co o ^ Q Q
g S 'a 1 -' ^ g
fc^<^i , t j
& 11
^ CN ON *O NO ON
CO CN CN CN CO H
CO-^-COCNCNCOCNCN
_C PQ C
03
S g
"S ^ .S
CO -( CN CO CO IO ON
co t^ NO co NO NO ON O
11 -S.S
CN CO CN CO CN CO *-H
^CNCOCOCNCNCN'* 1
s
f>
(A
"3
a
*o
^^S^SS^^^
ON .^ O
10 CN "- 1 i^ '""' CO
_jj d .CO
3 < 1 o Q ,|
_J b '^ c (uj^'bii'p.
^S^'-'^^^co
' ' o o o
oo^-S^-S^-S
^
NO CN ^ N-4 ^ CN JQ
^^q^S^^^^
1 1 1 1 1 1 1
s | | 1 | I I |
3
O
o
z
5
-H
-H O
O Ov
Ov
bi>
w
a
V
C/2
o
Sept 6 . .
76 982 440
o
o
o
o
o
" 13
49 515 000
7 500
o
o
o
o
" 20
63,144 000
o
o
o
o
o
" 27. .
45 854 000
218 400
o
o
o
o
Oct. 4. .
48 193 000
251 000
1 800 000
o
o
o
" 11
15,129 540
486 ' 000
o
o
o
o
" 18. .
17 367 000
25 000
o
o
o
Q
" 25
5 416 500
13 500
o
o
o
Q
Nov. 1 . .
25,325 500
2 000
o
o
o
o
" 8. .
14 564 000
o
3 600 000
25 000
o
Q
" 15
36 Oil 000
o
o
o
o
Q
" 22
" 29
23,494,000
73,719 000
o
38 500
o
o
Dec. 6 . .
56 400 500
o
o
o
o
o
" 13
148,740 000
o
1 800 000
o
o
o
" 20
116,344 800
o
o
247 200
6 000
o
" 27
67 965 800
o
o
69 600
o
o
Average
95,852 602
112 896
555 000
407 602
101 358
866 083
321
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Dinobryon
sertularia
var.
stipitatum
Eudorina
elegans
Euglena
acus
II
^
Euglena
oxyuris
Euglena
oxyuris*
Jan 11..
" 21
100
" 25. .
Feb. 3
" 8
" 15
" 22. .
Mar 1
8. .
" 15. .
3,600
40,000
40,000
" 22. .
800
" 29
Apr 5 . .
2,600
2,800
" 12
1,800
100
" 19
9,960
36,000
100
" 26
1,830,400
240,000
800
May 3 . .
4,883,200
240,000
3,200
" 10
24,608,000
48,800
" 17
28,800
32,800
90,000
180,000
" 24. .
1,000
" 31 ..
400
June 7
9,600
" 14. .
60,000
900 , 000
1,600
" 21 ..
30,400
2,400
" 28
4,000
1,800,000
July 5 . .
400
400
800
" 12
800
400
1,200
" 19
o
7,600
400
3,600,000
" 26. .
4,000
2,400
3,600,000
Aug. 2 ...
8,000
800
120,000
3,200
1,800,000
9
400
800
1,200
3,600,000
" 16
800
800
120,000
3,600,000
" 23. .
3,200
1,600
6,400
120,000
" 30. . .
2,400
800
3,200
4,500,000
Sept 6
40
1,600
900,000
10,400
5,400,000
" 13. .
500
1,500
3,600,000
" 20. ...
2,000
1,500
1,000
1,800,000
" 27
1,600
6,400
1,800,000
9,600
9,000,000
Oct. 4. .
1,500
3,600,000
1,000
2,700,000
" 11
1,000
1,800,000
500
1,800,000
" 18. ...
900,000
" 25. .
Nov. ' 1 . .
'
" ' 8
1,800,000
" 15
' 22. ..
1,000
1,800,000
" 29
120,000
Dec. 6 . .
o
" 13
500
" 20. ...
22,000
" 27..
603 911
14 362
375
214,807
963
960,769
322
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
O.eg
J>2
"3>-
^ B
Euglena
viridis*
Glenodinium
cinctum
Glenodinium
cinctum*
Gonium
pectorale
Lepocinclis
ovum
Lepocinclis
ovum*
Tan. 1 1 ....
80 000
o
" 21. .
o
o
100
Q
" 25. ..
o
o
Q
Feb 3
o
o
o
Q
8. .
o
o
Q
" 15..
o
Q
" 22
o
o
Mar 1 . .
o
o
Q
" 8
" 15
" 22. .
200
400
400
200
200,000
240,000
4 260 000
o
o
Q
" 29
240 000
o
Apr 5 . .
240 000
o
200
o
" 12. ..
120 000
200
Q
" 19
400
1,200
240 000
o
800
Q
" 26
May 3
3,200
360,000
120,000
22 400
o
Q
" 10.
" 17. .
120,000
3,600,000
o
90 000
200
800
o
Q
" 24. .
630,000
o
o
o
200
1 800 000
" 31
60,000
o
o
400
Q
June 7 ...
o
900 000
o
o
180 000
" 14
1,600
2,700,000
o
60 000
800
420 000
" 21
" 28. .
3,200
7,200,000
900 000
o
7,200,000
o
o
2,400
5 600
240,000
Q
July 5 . .
120,000
o
o
1 600
Q
" 12..
2 700 000
o
o
o
800
" 19. .
2 400
3 600 000
o
7 200 000
o
4 400
" 26. .. .
3,200
14 400 000
o
2 700 000
o
30 000
900 000
Aug 2 . .
1 600
7 200 000
20 000
12 600 000
o
50 400
9. .
4 800
7 200 000
400
25 200 000
o
6 400
" 16
5 400 000
o
5 400 000
o
800
720 000
" 23
4 800
4 500 000
o
o
o
14 400
" 30. .
8 000
2 700 000
800
* 900 000
800
43 200
Sept. 6 . .
800
3 600 000
900 000
o
11 200
240 000
" 13
" 20. .
1,000
3 000
1,800,000
500
120 000
o
1,000
5 000
" 27. .
6 400
1 800 000
o
3 200
8 000
Oct. 4
" 11 ..
6,300,000
120 000
o
o
o
2,500
2 000
1,800,000
Q
" 18
o
o
o
500
120 000
" 25..'
o
o
o
o
Q
Nov. 1 . .
o
o
o
o
500
o
8
o
o
o
o
o
Q
" 15
o
o
o
o
o
Q
" 22. .
o
o
o
o
o
O
Q
" 29
o
o
o
o
o
Q
120 000
Dec. 6. .
o
o
o
Q
" 13. ...
o
1 020 000
o
60 000
o
Q
Q
" 20
o
o
o
900 000
o
o
o
" 27. .
o
o
o
960 000
Q
Q
Average
8 653
1 571 731
452
1 360 192
526
3 719
401 538
323
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Mallomonas
producta
Pandorina
morum
Peridinium
tabulatum
Peridinium
tabulatum*
Phacus
longicauda
Phacus
pleuronectes
Platydorina
caudata
Pleodorina
calif arnica
Jan. 11
" 21. .
400
100
100
" 25
Feb 3 . .
100
8
400
" 15
" 22.
Mar 1 . .
400
8
" IS. .
600
" 22. .
" 29. .
200
600
Apr S . .
200
" 12. .
500
600
" 19
800
400
1,600
" 26.
48 , 000
3,200
May 3 . .
12,800
48 , 400
3,200
" 10
" 17. .
800
" 24. .
" 31
400
June 7 . .
835,200
8,000
120,000
200
" 14
28 800
60,000
900,000
8,800
800
" 21
" 28. .
28,800
28,800
40,800
9,600
2,400
8,800
1,200,000
79,200,000
8,800
4,800
800
July 5 . .
400
2,000
5,400,000
4,800
" 12.
800
18,800
10,800,000
3,200
400
400
" 19. .
12,000
49 , 600
86,400,000
3,200
400
400
120
" 26. .
63,200
66,800
15,300,000
6,800
800
400
Aug. 2
9. .
800
59,200
1,200
12,000
7,200
120,000
120,000
11,200
4,800
2,000
" 16. ..
3,200
8,000
" 23
2,400
6,400
1,800,000
4,800
800
.
60
" 30. .
3,200
6,400
8,000
1,600
Sept. 6. . . .
2,400
12,800
1,600
" 13
3 000
3,000
1,000
" 20. .
1
1,500
7,000
500
" 27..
1,600
100
4,800
35,200
4,800
Oct 4. .
7,000
" 11
" 18
S
500
1,500
1 ,000
" 25
500
Nov. 1 . .
500
8
" 15. .
1,000
1,000
" 22. .
" 29. ...
Dec. 6 . .
" 13
180,000
" 20. .
o
" 27. ..
Average. .
17 520
6 957
3 711
3 875 769
3,031
298
17
11
(22)
324
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Syncrypta
volvox
^
S to
?,5
<0
Synura
uvella*
Trachelomonas
acuminata
Trachelomonas
acuminata*
Trachelomonas
hispida
Trachelomonas
volvocina*
Jan. 11
" 21. .
100
5,600
3,600
3,800
" 25. .
7,740
387
Feb 3 .
1,600
4,600
480,000
8. .
800
800
60,000
" 15
10,800
3,600,000
28,800
" 22
Mar 1 . .
800
8,800
800
200,000
900,000
8
" 15. .
400
1,200
8,800
109,200
800
900,000
1,800,000
" 22. ..
221 ,600
60,000
10,800,000
" 29
Apr 5 . .
320,600
166,600
200
200
900,000
1,800,000
" 12
100
17,800
" 19
126,000
60,000
9,000 000
" 26 .
121 ,600
120,000
4,500,000
102 400
3 200
3 600 000
" 10. .
38,400
9,000 000
" 17. .
21 ,600
3,600,000
14,400,000
" 24
" 31 .
1,400
200
60,000
200
360,000
180,000
June 7
4,500,000
" 14
" 21 ..
1,600
800
120,000
7,200,000
7,200,000
147 600 000
" 28. .
800
6,300,000
38 700,000
Tuly 5 .
400
1 800 000
1 800 000
" 12! !
1 ,200
800
900,000
10 800 000
" 19
800
3,600,000
86,400,000
" 26
2 000
3 600 000
9 200
42 300 000
Auff 2 . .
12,800
600,000
800
18 000 000
.? g
800
3,600,000
252,000,000
" 16..
4,000
3,600,000
1 ,600
93,600,000
" 23. .
3 200
1 800 000
800
65 700 000
" 30. .
8 800
1 800 000
1 ,600
18 000 000
Sept. 6
" 13..
4,000
5 , 400 , 000
800
1 000
16,200,000
6 300 000
" 20. .
4,000
1 500
1 800,000
" 27
Oct. 4. .
1,600
4,800
500
3,600,000
1 800 000
1,600
9,000,000
11 700 000
" 11
120 000
1 ,800 000
" 18
" 25
Nov. 1 . .
500
2,000
900,000
120,000
2,700,000
5,400,000
1,800,000
8 .
1 000
16 000
1 800 000
" 15..
9 000
" 22
94,000
1 800 000
5 400,000
" 29
4,500
1 ,999,500
1 ,320,000
3,600,000
Dec. 6 . .
13,500
1 693 500
2 280 000
900,000
" 13
" 20. .
2,000
6 200
78,000
2 764 800
2,760,000
900 000
o
500
o
2,400,000
" 27... .
800
395 200
300 000
o
2 700 000
Average. .
625
179.138
150.000
873
t .094.615
1.251
17.672.692
325
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Total
Rhizopoda
Arcella
discoides
A rcella
vulgaris
Centropyxis
aculeata
Centropyxis
aculeata var.
ecornis
Cochliopodium
bilimbosum
Cyphoderia
margaritacea
Difflugia
acummata
Jan 11..
440,500
100
" 21
32,800
100
200
100
" 25
66 338
387
387
387
1,161
20,898
774
1,935
Feb 3 . .
122,900
1,300
8
4 880
3,200
" 15 .
34 880
800
800
800
80
" 22. .
141 ,524
632
25,272
12,636
9,477
3,159
Mar 1
11 200
400
400
400
400
8. .
11 ,720
400
400
1,200
400
40
" 15. .
7,600
600
400
" 22
" 29
4,800
61 400
400
400
200
Apr 5 . . .
700
100
100
100
" 12
3 520
300
200
20
100
" 19
" 26 .
7,300
6,720
400
400
400
May 3 . .
26,000
400
" 10
" 17
" 24. .
49,800
23,800
9,320
2,400
600
1,600
800
200
400
80
" 31
June 7 . .
8,920
23 600
400
800
200
200
400
200
3,200
" 14. .
21,600
1,600
800
" 21
21 600
1 600
800
800
" 28. .
37 000
800
800
100
July 5
19 360
1 200
400
400
400
" 12
26 000
800
800
200
o
1 600
1 200
" 19. .
28 800
800
400
400
400
" 26. .
4 800
400
400
400
Aug 2
16 800
800
4 800
1 600
9. .
7 280
1 600
400
1,600
40
40
" 16. .
24 060
2 400
800
800
800
" 23
36 800
800
5 600
800
" 30
Sept 6 ... .
23,200
20 800
800
800
5,600
800
800
1,600
3,200
1 600
" 13
28,000
500
500
6,000
1 ,000
" 20
" 27. .
19,000
59 200
500
1 600
500
1 600
500
500
1,000
8,000
1,500
o
500
3 200
Oct. 4. .
912,580
40
40
500
500
" 11
9 000
1 000
" 18. .
10 000
500
2 000
1 000
" 25.. .
25,060
1,000
1,500
1,000
1,000
500
500
500
Nov 1.
32 060
500
1 000
1 000
2 000
500
500
8. .
37 060
1 000
1,000
1 ,000
1,000
1 000
" 15
42,000
1,000
5,000
4,000
" 22
190 400
2 000
4 000
6 000
o
" 29. .
3 400
o
500
500
o
Dec 6.
121 000
o
1 000
o
" 13. .
600
600
" 20
1 040
40
1 ,000
" 27. .
220
o
55 364
465
1 098
570
604
1 284
198
315
326
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Difflugia
globulosa
Difflugia
lobostoma
Difflugia
pyrijormis
Total
Heliozoa
Nuclearia
delicatula
a
ll
SG
EH
Amphileptus
spp.
Carchesium
lachmanni
Jan 11..
100
100
593,420
26 500
" 21
400
200
197,100
37 000
" 25. .
9,675
7 353
190 017
13 545
45 666
Feb. 3 . .
500
100
1,246,300
1 600
54 700
8
" 15. .
800
9,200
80
2,000
629,680
1 016 000
800
4 400
197,600
164 800
" 22. .
6,318
632
518,954
50 544
Mar 1 . .
4,000
800
400
1 773 360
400
46 400
8. .
2,800
800
40
492 920
800
54 800
" 15. ..
2,600
1,400
200
324,200
200
89 600
" 22
1 600
800
2 000
267 800
400
22 000
" 29. .
200
400
241 420
400
10 200
Apr 5
100
500
241 440
o
3 100
" 12
" 19. .
1,000
1,600
800
1,200
100
100
1,342,500
1,340 800
300
2,400
13 200
" 26. ...
3,200
3,200
7,710 400
99*200
May 3 . .
22,400
3 200
17 404 800
83 200
" 10
30,400
3,200
18 260 800
6 400
" 17
800
800
21,654,400
1 600
" 24. .
3 640
200
200
990 800
o
200
" 31. .
3,840
400
2 282 400
o
600
June 7
" 14. .
9,600
5 600
8,000
1 600
200
800
o
7 , 800 , 000
6 480 000
o
o
" 21..
5,600
800
3 200
3 200
12 010 400
o
o
" 28. ...
14,400
2,400
100
6 491 200
o
9 goo
July 5 . .
8 800
2 000
160
495 640
o
o
" 12..
10,000
2 400
800
400
400
3 900 920
o
400
" 19
12 800
2 000
2 000
2 000
721 640
o
400
" 26. .
2 400
400
14 400
14 400
487 'OOO
o
o
Aug. 2 . .
5,600
800
17 600
17 600
4 474 400
o
o
" 9
" 16. .
2,800
8 000
400
800
3 200
78,400
13 600
78,400
13 600
69,000,200
253 600
o
o
" 23. .
12 800
2 400
20 800
20 800
728 000
o
1 600
" 30. .
6 400
800
800
7 200
7 200
122 400
o
o
Sept 6 . .
5 600
o
800
4 800
4 800
120 001 640
o
800
" 13. .
11 500
500
500
500
1 451 120
o
9 000
" 20. . .
8 500
500
18 000
18 000
1 923 'SOO
o
2 500
" 27
25,600
1 600
1 600
65 000
65 600
851 400
o
o
Oct. 4. .
8 000
500
500
720 000
o
o
" 11. ...
2 500
1 000
843 540
o
3 500
" 18
" 25
Nov. 1 . .
8
" 15..
2,000
15,000
15,000
5,000
17 000
1,000
1,000
2,000
2 000
60
60
2,000
500
500
o
500
500
o
1,744,000
1,334,000
985,560
965,000
488 100
500
500
2,000
1 000
5,000
35,000
31,000
22,000
28 000
" 22...
48 000
8 000
400
750 640
4 000
108 000
" 29
200
200
721 "00
o
47 500
Dec. 6 . .
o
o
720 580
40
7 000
" 13. ...
o
1 573 800
100
16 400
" 20. .
o
o
o
o
487 120
o
16 600
" 27. . .
200
o
o
o
490 600
200
28 000
Average
7 194
1 158
368
4 871
4 760
15 812 346
630
26 546
327
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Codonella
cratera
Halteria
grandinella*
Stentor
cceruleus
Tintinnidium
fluviatile
Total
Suctoria
Metacineta
mystacina
2
11
^
1
Jan 11
300
80,000
300
6,580
" 21
" 25. .
300
58,437
40,000
28,800
11,997
100
49,240
126^03
Feb 3
5,900
1,000
1 1 , 496
" is! !
8,000
5,200
800
800
14,160
31,040
" 22
15,795
720,000
48,649
Mar 1 . .
10,000
1,600
1,600
20,400
400
8,400
520
1,600
1,600
29,200
800
" 15
33,200
1,000
400
400
103,940
400
" 22
41 ,600
60,000
80
1,200
1,200
185,520
400
" 29
Apr 5
30,400
20,500
- 20
300
200
100
200
115,880
84,820
5,020
1,800
" 12 .
20,100
900,000
200
100
54,540
" 19
" 26
453,600
614,400
400
12,800
749,000
2,892,360
4,800
May 3 . .
" 10. .
736,000
78,400
720,000
24,000
5,247,800
2,663,400
200
" 17
72 000
10,400
800
800
1,465,500
800
" 24
74,200
400
196,020
3,200
" 31
June 7
61,200
1 499 200
60 000
400
14,400
200
200
180,760
903,000
18,800
392,000
" 14 .
532 ,800
104,000
639,600
1,600
" 21 ..
195, ?00
74,400
800
2,601,200
3,200
" 28
45,600
3,600,000
33,600
1,118,400
July 5
" 12. .
13,600
35,600
2 700,000
4,800
5,600
7,200
400
7,200
400
153,000
184,500
800
" 19
24,000
2,800
400
400
946,080
" 26
2,000
120,000
3,600
370,200
Aug 2 . .
23,200
1 800,000
95,200
1,294,240
9
" 16
8,400
20,000
4,800
8,800
782,720
935,380
" 23. .
26,400
5,600
1,600
1,600
696,180
1,600
" 30
51,200
800
435,080
1,600
Sept 6 . .
13,600
40
422,840
" 13. .
49,000
120
2,000
197,960
" 20
34,500
20,000
500
475,860
1,000
" 27
92 800
200
22,400
1,792,700
14,400
Oct. "4. .
23,000
1,500
105,020
2,580
" 11
23 000
40
500
122,000
2,000
" 18. .
47,000
900,000
1,000
40
40
159,200
" 25. .
23,000
1,048,620
Nov. 1 . .
8. .
12,500
70 000
60
156,300
147,780
" 15. ...
35,000
100
180,600
" 22
" 29. .
2,000
2 000
128,400
66,000
Dec 6
40
64,280
" 13. .
300
1 080 000
159,740
" 20
200
120,000
191,320
" 27.
200
120 000
o
50,540
200
Average
101,024
255,769
882
22,590
332
301
592,416
328
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Conochilus
dossuarius
Conochilus
unicornis
Total
Bdelloida
Philodina
megalotrocha
Rotifer
neptunius
11
'S 2
CS
.|
"*.
fr;
Anuraa
aculeata
Jan. 11 ..
400
400
6,180
" 21
45 100
44 500
4 040
" 25. .
90,171
89,379
35 271
Feb 3
3 800
3 800
7 696
8
" 15. .
6,800
18,000
6,800
27,000
7,360
12 240
" 22
25 272
25 272
23 377
Mar 1 . .
1,600
400
800
18 320
8
400
4,040
40
4,000
23,960
400
" 15
400
22 160
80
400
19 200
80 980
40
" 22
" 29. ..
400
20
10,440
1,620
40
20
10,400
1,600
174,680
109,240
400
200
Apr. 5 . .
" 12. ..
1,100
960
60
1,100
800
81,920
53,480
600
600
" 19
400
3 300
400
100
16 000
745 300
2 000
" 26. .
3,200
4,640
640
3,200
2 889 720
3 200
May 3
16 000
12 800
5 231 800
22 400
" 10
" 17
" 24. . .
200
800
3,200
14,400
20,800
1,040
80
1,600
6,400
520
11,200
10,400
400
2,647,200
1,438,300
191 780
35,600
22,400
4 000
" 31
o
18 600
880
80
200
600
161 080
1 400
June 7 ....
" 14
392,000
1,600
800
600
800
400
200
507,000
637,400
1,600
800
" 21
3 200
1 100
300
800
2 593 600
" 28. .
1 900
800
300
1 112 500
July 5
800
2 480
80
1 600
800
146 920
o
" 12
" 19. .
4,800
2 760
400
1 600
2,000
360
2,400
800
178,100
933 320
" 26. ..
120
60
268,480
Aug 2
1 400
560
40
1 260 840
o
9. .
1 200
12 000
775 920
o
" 16. ..
4 120
120
4 000
907 260
o
" 23
1,600
5,720
60
5,600
671 ,260
" 30
1 600
o
4 080
80
2 400
415 000
o
Sept. 6 ...
9 640
2 400
40
4 800
413 200
o
" 13
21,000
500
1,500
17,500
171,960
" 20. .
1 000
o
6 000
1 500
500
3 000
460 360
o
" 27. .
14 400
o
13 300
8 000
300
4 800
1 744 200
o
Oct 4
2 500
o
2 280
2 000
120
160
97 160
o
" 11 ..
2 000
o
2 000
1 000
500
500
115 SOO
o
" 18. .
o
540
40
188 160
o
" 25
3,500
1 000
2,500
1,045 120
o
Nov. 1 . .
o
3 060
60
3 000
152 680
o
8. . .
1 180
120
60
1 000
146 600
" 15.
o
100
100
180 500
o
" 22. .
o
400
400
126 000
o
" 29..
o
66 000
o
Dec. 6 . .
o
20
20
64 260
o
" 13..
o
600
600
159 140
o
" 20. ...
191 320
" 27..
200
o
20
o
20
o
50 120
20
Average
517
8 108
405 983
351
425
6 688
571 611
1 839
329
TAB-LE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Anurcea
cochlearis
type and var.
macracantha
Anurcea
cochlearis
var. stipitata
Anurcea
cochlearis
var. tecta
Total
Anurcea
cochlearis
Total eggs
Anurcea
cochlearis
Atmrcea
hypelasma
Total eggs
Anurcea
hypelasma
Jan 11..
100
100
" 21
300
300
100
" 25
387
1,661
2,048
Feb 3 . .
500
100
600
300
8
80
80
" IS. .
80
80
" 22. .
Mar 1 .
400
80
480
8. .
800
1,200
2,000
1,600
" IS. . .
2,200
600
2,800
1,400
" 22
3 200
800
4,000
2,800
" 29.
2,600
600
3,200
200
Apr. 5 . .
1,700
400
2,100
600
" 12
1 800
400
2,200
800
" 19. .
12,400
2,800
15,200
8,800
400
" 26. .
12,800
121,000
4,000
137,800
57,800
222 400
745,600
54,400
1,022,400
552,200
" 10. .
134,400
790,400
220,800
1,145,600
643,200
" 17. .
91 ,200
295,600
48 , 000
434,800
160,000
" 24
1 000
18 400
1,800
21,200
7,200
" 31 .
1 400
9,200
600
11,200
3,400
June 7 ....
32,000
32,000
3,200
" 14
28,000
1,600
29,600
7,800
2,400
" 21 ..
150,400
222,400
372,800
148,800
9,600
8,800
" 28. .
48,800
117,600
166,400
20,800
7,200
4,000
July 5. .
2,800
7,200
10 000
1,600
800
400
" 12. .
2,000
8,000
10,000
4,000
1,200
" 19. .
2,000
15,200
17,200
5,600
4,000
" 26
1 200
1 200
Aug 2 . .
4,800
2,400
9. .
2,000
4,000
" 16
16 000
8 800
" 23. .
9 600
3,200
" 30. .
800
800
Sept 6
" 13. .
500
500
1 ,000
" 20. .
3,500
8,500
12,000
6,000
4,000
1,000
" 27
19,200
35,200
54,400
16,000
43,200
54,400
Oct 4. .
4,000
2,000
4,000
500
2,000
500
" 11 ...
7,000
2,000
9,000
4,500
500
" 18
o
17 500
7 000
24 500
10 500
3 500
2,500
" 25. .
500
9,000
19,000
28,500
7,000
13,500
5,000
Nov. 1 . .
500
1,000
1,500
,0
500
1 ,000
8
" 15. .
" 22. ...
" 29
500
1,000
8,500
10,000
500
Dec. 6 . .
1 , 000
1,020
2,200
20
" 13. .
500
1 700
5 100
7 300
1 800
" 20. ..
3 600
1 ,600
5,200
2,600
" ' 27
200
200
Average
9,421
44 540
15,432
69,165
32,358
2,390
1,917
330
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Asplanchna
brightwellii
Asplanchna
priodonta
Brachionus
angularis
Brachionus
angularis
var. bidens
Total
Brachionus
angularis
Total eggs
Brachionus
angularis
Brachionus
bakeri
var.
brevispinus
Jan 1 1 .
" 21 ..
100
" 25
387
387
Feb 3 . .
8
" IS
o
" 22. .
Mar 1
80
8. .
" IS
" 22
40
" 29
Apr 5
20
100
100
" 12
" 19. .
" 26. ..
May 3 . .
16 000
" 10. .
20,800
3,200
1,600
1,600
" 17
11 ,200
14,400
800
800
800
" 24
" 31 ..
400
200
2,120
2,000
1,400
200
200
1,400
June 7 . .
" 14
" 21 . .
200
1 ,100
1,100
4,800
4,000
70,400
4,800
4,000
70,400
1,600
24,800
" 28
100
544,000
544,000
128,800
July 5 .
160
29,200
400
29,600
1,600
" 12. .
51,200
51,200
13,200
400
" 19
280
300,800
34,800
335,600
72,800
" 26
17 900
6 400
10,400
16,800
1,200
Aug. 2 ...
23,200
10,400
93,600
103,200
12,000
9
80
229,200
64,800
292,600
105,600
400
" 16
800
272,800
80,800
353,600
116,000
" 23. .
4,000
77,600
138,400
216,000
42,400
" 30
2,400
28,800
86,400
115,200
28,000
2,400
Sept 6. .
80,000
83,200
163,200
35,200
400
" 13. .
60
27,000
10,000
36,500
18,000
2,000
" 20
1,140
60
87,500
27,500
115,000
43,000
" 27
6,400
409 , 600
84,800
494,400
41,600
1,600
Oct. 4. .
500
19,000
9,000
28,000
2,000
" 11
1,000
8,000
1,000
9,000
2,000
" 18
8,000
500
8,500
2,500
" 25..
60
11,500
11,500
5,500
Nov. 1 . .
1,000
,
1,000
8. ...
" IS...
100
100
" 22
" 29
Dec. 6 . .
20
20
" 13
" 20. .
400
400
" 27
2 079
441
43 946
13,973
57,919
13,242
139
331
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Brachionus
bakeri
var. clunior-
bicularis
Brachionus
bakeri
var. melhemi
Brachionus
bakeri
var. obesus
Brachionus
bakeri
var.rhenanus
Brachionus
bakeri
var.
tuberculus
Total
Brachionus
bakeri
II
*|'l
d g^
ocqj
H
Jan 11..
" 21 .
" 25. .
Feb 3
8
" IS. .
" 22
Mar 1 .
8. .
" IS
" 22
40
" 29
Apr 5 . .
" 12
" 19 .
" 26. .
May 3
" 10 .
" 17. .
" 24
" 31
40
40
" 14
" 21
" 28. .
800
100
900
Tulv 5
400
400
800
400
' 12!
60
1,200
1,660
60
" 19. ...
920
1,200
40
2,160
2,520
" 26
Aug 2 . .
9. ...
400
40
840
800
" 16
800
1,600
2,400
5,600
" 23
" 30. .
800
800
800
800
5,600
2,400
Sept. 6
800
800
1,600
4,000
7,600
5,600
" 13
500
2,000
4,500
4,000
" 20. .
500
500
500
" 27..
1,600
3,200
Oct 4 .
40
" 11 ..
" 18. .
40
40
" 25
500
500
Nov 1 . . .
60
60
8
" IS.
" 22. .
" 29
Dec. 6. .
" 13
" 20. .
" 27. ..
Average
90
49
41
118
155
592
420
332
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Brachionus
budapesti-
nensis
Brachionus
militaris
Brachionus
mollis
Brachionus
pala
Brachionus
pala
var.
amphiceros
Brachionus
pala
var. dorcas
Brachionus
pala var.
dorcas forma
spinosus
Total
Brachionus
pala
Jan. 1 1 ...
" 21. .
o
o
o
20
100
20
o
20
o
o
60
100
" 25. .
o
o
o
o
387
387
Feb. 3 . .
o
o
o
o
200
o
200
8. .
o
o
o
o
o
o
o
o
" IS. . .
o
o
o
o
o
o
o
" 22
o
o
o
o
o
o
o
Mar 1 . .
o
o
80
o
o
o
80
8
o
o
o
80
o
80
" 15
" 22. .
o
o
o
160
o
o
360
1 720
o
520
1 720
" 29
o
o
o
200
o
140
o
340
Apr. 5 . .
" 12. ..
o
o
o
200
20
120
100
160
o
120
480
" 19
o
o
2 800
1 200
800
o
4 goo
" 26. .
o
o
o
57 920
97 600
4 000
o
159 520
May 3 . .
o
o
32 000
419 200
o
o
451 200
" 10 .
o
o
o
19 200
57 600
o
o
76 800
" 17. .
o
o
o
5 600
69 600
800
1 700
77 700
" 24. .
o
200
o
80
200
o
o
280
" 31
June 7 . .
o
o
o
200
o
o
o
200
" 14
o
o
o
o
o
o
o
1 000
" 21
" 28.
4 000
o
o
800
o
200
o
o
o
o
July 5 . .
3 600
40
o
40
o
o
o
40
" 12
10 000
120
o
o
o
o
o
" 19. .
85 600
80
o
800
5 600
o
o
6 400
" 26. .
3 200
o
o
o
120
o
o
120
Aug. 2 . .
9 600
40
o
6 400
o
o
6 400
9..
11 200
o
200
1 200
2 000
o
o
3 200
" 16. .
20 000
o
800
800
37 600
o
o
38 400
" 23
8 000
800
o
o
35 200
o
o
35 200
" 30
7 200
3 200
800
800
32 000
o
o
32 800
Sept. 6. .
7 200
1 600
o
o
19 200
o
o
19 200
" 13
1 500
o
o
500
4 000
o
o
4 500
" 20
2 000
500
500
9 000
o
o
9 500
" 27. .
44 800
1 600
4 800
4 800
78 400
o
o
83 200
Oct. 4 . .
80
o
40
1 000
o
o
1 040
" 11
500
o
o
o
500
" 18
" 25..
1,000
o
o
500
3 500
80
5 000
o
o
580
8 500
Nov. 1 . .
8
" 15..
o
o
o
1 000
180
100
o
o
180
1 100
" 22...
o
o
o
400
400
o
o
800
' 29
Dec. 6. .
o
o
o
1,000
320
500
1 160
20
o
1,500
1 500
" 13
" 20. ...
" 27..
o
o
o
2,400
1,200
400
3,200
606
200
40
o
5,600
1,800
640
Average
4 211
147
137
2 693
17 071
170
33
19 969
333
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Total eggs
Brachionus
pala
Brachionus
urceolaris
Brachionus
urceolaris
var.
bursarius
Brachionus
urceolaris
var. rubens
Total
Brachionus
urceolaris
Total eggs
Brachionus
urceolaris
Brachionus
variabilis
Brachionus
free winter
eggs
Jan. 11..
" 21. .
100
100
40
40
100
" 25..
1,161
1,548
Feb 3
100
8. .
" 15. ..
800
2,400
" 22
632
3,791
Mar 1 . .
480
80
80
480
8. ...
160
840
" 15
1,000
160
160
400
" 22
2,920
2,000
2,000
400
440
" 29.
420
1,800
1,800
1,200
20
Apr. 5 . .
20
700
700
400
120
" 12
240
140
140
60
200
" 19
5,200
400
400
" 26. .
324,280
6,400
6,400
May 3
661,200
41,600
" 10.
118,400
9,600
" 17. .
101,700
800
" 24.. .
1,040
200
80
" 31
400
June 7 . .
" 14. ...
400
400
" 21
100
800
800
" 28. .
100
100
100
100
July 5 . .
40
40
" 12 .
400
1,200
" 19. .
400
40
200
240
400
" 26. ..
800
400
400
1,200
Aug 2
1,200
o"
9. .
8,400
800
800
" 16. ..
5,600
800
800
800
" 23
" 30
14,400
12,000
2,400
2,400
800
4,800
Sept. 6. . .
22,400
5,600
5,600
800
" 13
" 20
" 27. .
3,000
5,500
32,000
500
500
500
. 1,600
Oct 4
500
500
" 11
500
" 18. .
500
.
540
" 25
4,500
500
Nov 1 . .
2,000
8. .
120
2,000
" 15
" 22. .
1,200
2,400
1,000
2,000
" 29. .
1,500
500
Dec 6 . .
860
100
100
" 13. ..
10,600
600
600
" 20
1,800
40
40
" 27. .
100
240
240
80
1,621
25 974
18
206
244
468
56
4
1,685
334
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Mastigocerca
carinata
Metopidia
solidus
Monostyla
bulla
Monostyla
lunaris
Notholca
striata var.
acuminata
Polyarthra
platyptera
Polyartha platyptera
Male eggs
Free
Carried
Jan 11
400
19,200
11,200
2,000
1,600
7,200
4,000
15,200
4,000
5,600
5,600
800
2,500
1,500
4,800
1,000
60
400
400
100
400
800
800
800
200
100
100
400
200
200
o
800
800
100
800
400
500
24
240
80
800
1,200
6,400
10,800
200
300
40
1,000
1,200
11,997
3,200
2,000
1,600
6,318
3,200
5,200
22,200
37,600
40,400
42,800
26,700
148,200
696,000
582,400
137,600
195,200
52,200
52,400
304,000
432,800
241,600
56,800
6,400
21,600
89,200
86,400
288,000
55,200
84,800
96,000
51,200
4,000
31,000
72,500
238,400
24,500
47,500
27,000
37,500
500
1,000
2,000
6,000
1,000
6,020
42,100
63,400
19,200
1,300
900
8,800
150,400
4,800
12,000
200
1,600
5,000
2,000
100
1,600
2,600
2,800
1,900
1,600
53,800
19,200
2,400
1,600
500
1,500
2,000
160
100
200
" 21
" 25. .
Feb 3
8.
" 15..
" 22
Mar 1 . .
8
" 15
" 22. .
" 29. .
Apr 5 .
" 12. .
19
" 26
May 3 . .
" 10
" 17
" 24. .
" 31
June 7 . .
" 14. .
" 21 .
" 28. .
July 5 . .
" 12
" 19. .
" 26. .
Auf? 2
" 9. .
" 16. .
" 23
" 30. .
Sept. 6 . .
" 13
" 20. .
" 27. .
Oct. 4. ...
" 11. .
" 18. .
" 25
Nov. 1 . .
8
" 15
" 22. .
" 29
Dec 6. .
" 13. .
" 20
" 27. .
Average .
1,674
67
50
.57
388
86,674
3,598 ;
1,768
335
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter^paper collections.)
1898
Polyarthra
platyptera
Winter eggs
2 2
It
"
If 8
*"
Pterodina
patina
Rattulus
tigris
Schizocerca
diversicornis
Synchceta
pectinata
Synchceta
stylata
Total free
winter eggs
Synchceta
Free
Carried
Jan 11..
100
800
200
200
100
1,600
22,400
51,200
11,200
2,400
400
400
800
800
800
400
400
800
800
800
1,000
1,000
1,600
1,000
500
1,000
1,000
100
100
1,600
2,200
1 ,000
8,127
1,700
2,800
3,200
6,318
3,200
4,000
17,800
26,400
11,400
10,400
8,700
104,200
502,400
316,800
72,000
120,000
28,800
10,600
96,000
119,200
154,400
16,000
7,200
13,600
24,000
53,600
295,200
84,800
108,000
63,200
47,200
8,800
20,000
103,000
86,400
15,000
5,500
14,000
17,000
4,500
2,000
2,000
6,000
3,000
9,160
29,300
52,000
1 1 , 000
200
40
1,600
100
100
400
40
400
100
300
200
3,200
200
40
400
800
400
800
800
1,600
1,000
800
800
800
.
72
632
400
1,600
800
200
600
3,200
800
112,000
800
400
20,800
400
12,000
4,800
1,600
3,200
800
1,000
4,500
30,400
500
500
500
20
2,500
400
2, \20
300
4,257
1,000
1,200
800
6,400
4,800
15,200
58,000
47,000
2 1 , 000
11,500
368,000
954,400
1,139,000
233,600
206,400
60,480
61,600
48,000
19,200
795,200
22,400
22,800
9,600
64,800
8,000
170,400
52,000
18,400
24,800
1,600
14,000
27,000
265,600
5,000
27,000
77,000
824,500
110,500
97,000
110,000
38,000
39,000
42 , 500
55,720
59,200
17,640
100
1,200
160
6,400
9,600
6,400
800
800
3,200
400
800
800
60
.
500
500
100
60
" 21
" 25 .
Feb 3 . .
8
" 15. .
" 22. .
Mar 1
8. .
" 15. .
" 22. .
" 29
Apr 5 . .
" 12. ..
" 19
26"""
May 3 . . .
" 10
" 17. .
" 24. .
" 31 ..
June 7 . .
" 14. .
" 21 ...
" 28. .
July 5 . .
" 12. ...
" 19
" 26. .
Aug. 2 ....
9
" 16. .
" 23. .
" 30. .
Sept 6 . .
" 13
" 20...
" 27
Oct. 4. .
" 11 ...
" 18. ...
" 25..
Nov. 1 . .
8. .
" 15. .
" 22. ..
" 29 .
Dec. 6. .
" 13
" 20. .
" 27. ..
Average . .
1,994
34
52,560
37
207
46
3,950
120,391
611-
336
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Total eggs
Synchceta
Triarthra
longiseta
Triarthra
longiseta
eggs
Pedalion
mirum
Total
Entomostraca
Total
Ostracoda
Total
Cladocera
Jan 11
500
700
100
" 21..
200
1 380
440
" 25. ..
6,579
4 788
462
Feb 3 . .
300
216
24
8. .
400
320
" IS
5,600
1 200
80
" 22
Mar 1 . .
1,200
80
3,285
804
160
8
800
3 080
40
80
" IS. .
3 960
12 880
40
560
" 22. ..
4 000
40
40
19 440
320
800
" 29
3,200
100
22 180
160
360
Apr 5 . .
1,100
300
200
o
34 560
200
360
" 12
1,000
200
100
o
28 060
320
320
" 19
" 26. .
84,000
38 400
400
3 200
o
34,200
56 800
200
800
400
1 920
May 3 . .
278,400
9,600
o
204 800
2 800
" 10
91 600
38 400
o
o
235 400
400
5 600
" 17. .
56 800
17 600
3 200
o
182 300
1 600
8 500
" 24
9,400
600
200
o
167 080
400
24 080
" 31
12 000
1 000
o
162 800
440
51 480
June 7 . .
11 200
200
o
438 800
1 600
136 000
" 14
13,600
o
211 400
400
29 200
" 21
257 600
800
100
83 100
200
2 300
" 28. .
51 200
800
500
45 600
400
10 100
July S
" 12
" 19. .
47,200
16,800
34 800
400
1,600
4 000
400
1 , 600
1,200
9 200
4,920
1,620
14 040
440
60
o
640
360
1 240
" 26
4 800
28 000
1 600
99 600
23 000
o
9 960
Aug. 2 . .
178,400
18 400
1 ,600
30 400
22 160
40
10 520
" 9
20 000
4 400
1 200
4 000
4 120
o
1 080
" 16..
10 460
3 200
22 400
4*500
800
1 320
" 23
35,200
4 000
17 600
17 340
180
1 820
." 30.
7 200
6 400
o
14*400
27 080
o
4 040
Sept 6 ....
9 080
720
" 13....'..
9,500
1 000
5 000
24 720
SOO
3 420
" 20. .
17 500
500
o
5 SOO
21 880
o
1 560
" 27..
38 500
14 400
3 200
19 200
99*300
o
1 100
Oct. 4 . .
1 500
500
2 000
33 880
1 320
" 11 ..
2 000
o
2 000
34 060
o
2 000
" 18. .
10 500
1 500
500
500
25 640
2 120
" 25
78 000
500
o
o
26*020
1 020
Nov. 1 . .
29 000
o
o
60
8 600
o
120
8
41 060
IS 080
" IS
60,000
13 100
100
" 22
66,000
13 920
320
2 800
" 29..
500
500
500
o
6 180
o
80
Dec. 6. ...
o
9 740
260
" 13. .
800
o
o
o
21 740
o
240
" 20
2 600
o
o
2 440
400
" 27
400
. 40
6 800
280
Average
31 620
3 147
255
4 524
47 042
191
6 241
337
TABLE I ^continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
!
Bosmina
longirostris
Ceriodaphnia
scitula
Chydorus
spharicus
Daphnia
cucullata
S 2
II
tl
a-
Diaphanosoma
brachyurum
Moina
micrura
Jan 11
100
" 21 .
240
200
" 25
154
308
O,
Feb 3
24
8. .
" 15. .
" 22
Mar 1 .
80
80
8. .
40
40
" 15
120
440
" 22
280
480
" 29 .
20
20
240
20
Apr 5 . .
20
100
40
200
" 12
20
60
20
200
20
" 19
200
100
o
" 26
800
320
800
May 3 . .
2,800
" 10
3,600
400
600
600
" 17
200
3,500
400
3,300
300
" 24
" 31. .
480
40
5,920
33,920
2,960
8,720
7,880
5,040
440
1,000
160
720
40
200
62,800
55,800
600
3,400
11,600
" 14
6,000
10,600
200
2,400
9 200
" 21 .
100
1,500
400
100
200
" 28. .
200
700
100
Tulv 5
200
40
400
' 12::
180
60
120
" 19. .
160
40
1 040
" 26
180
120
o
o
o
8 580
1 080
Aug 2 . .
40
6 960
3 520
9. .
360
360
" 16
60
1 260
" 23
o
o
1 020
900
" 30
40
2 520
1 440
Sept 6
40
o
o
o
240
440
" 13. ...
60
1,800
1 560
" 20
60
60
o
960
480
" 27. .
100
o
400
500
Oct 4
40
400
o
880
120
" 11 .
920
600
o
400
40
" 18. .
1 360
80
80
o
560
" 25..
840
120
60
o
Nov 1
60
o
60
o
8. .
o
o
" 15..
100
o
o
" 22
o
o
o
o
o
" 29.
o
80
o
o
Dec. 6. .
40
20
200
o
" 13
' 20. .
140
120
40
o
220
160
o
o
" 27. .
o
280
o
o
o
Average
36
2,441
1,539
422
181
417
479
261
338
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
Total
Copepoda
Canthocamptus
spp.
Cyclops
albidus
B
ft. =0
O S
ft
^
-2
^t3
*
Cyclops
prasinus
Cyclops
virdis var.
brevispinosus
$
>2
" *
IH
"G-fe s
$**
Jan 11..
600
160
o
o
" 21
940
40
160
o
40
" 25
4 326
77
308
o
308
Feb 3 ...
192
48
o
o
8
320
160
o
o
" 15. .
1 120
80
o
80
o
" 22. .
3 285
o
o
o
Mar 1
644
o
o
o
8. .
2 960
40
120
o
o
" 15
12 280
40
400
o
80
120
" 22
" 29. .
18,320
21 660
200
100
20
80
60
o
20
80
o
Apr. 5 . .
34 000
200
20
40
o
" 12
27 420
1 120
o
o
40
" 19. .
33 600
200
200
o
500
" 26
54 080
1 600
2 880
o
o
4 160
May 3.
202 000
400
8 000
400
o
1 200
" 10. .
229 400
600
5 200
o
o
600
2 200
" 17
172 200
800
200
600
o
o
3 300
" 24
" 31. .
142,600
110 880
80
920
200
320
80
o
1,080
400
1,640
640
301 200
400
2 600
4 000
" 14
181,800
200
200
800
4 400
" 21
" 28. .
80,600
35 100
o
o
o
o
o
400
200
July S
3 840
o
o
o
o
o
120
" 12..
1 200
o
o
o
o
180
" 19. .
12 800
o
o
40
o
40
240
" 26
13 040
o
o
120
o
120
300
AUK 2 . .
11 600
o
o
o
40
AUK. *.-
3 040
o
o
80
o
160
" 16. .
2 380
o
o
o
180
" 23
15,340
o
120
660
" 30. .
23 040
o
o
o
440
o
480
Sept 6 ...
8 360
o
40
80
" 13
20,800
120
240
" 20
20,320
120
60
360
" 27. .
98 200
100
700
o
300
o
200
700
Oct. 4. .
32,560
120
320
200
400
" 11
32,060
200
80
120
" 18. .
23 520
o
280
o
o
40
40
" 25..
25 000
o
60
60
120
240
Nov 1 . .
8 480
o
120
o
o
60
8. .
15 080
o
o
120
" 15
13 000
200
200
" 22
10 880
480
80
160
" 29. .
6 100
80
o
4
o
o
Dec. 6 ...
9 480
20
20
" 13. .
21 500
o
40
40
" 20. . .
2 040
40
o
160
40
" 27
6 520
120
40
Average
40,609
78
113
373
49
2
124
539
339
TABLE I continued.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1908.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
1898
&|
II
S*
Copepodan
nauplii
ll
!
?*
Diaptomus
siciloides
a a
a*
Total
Nematodes
Total
Oligochsetes
Chironomus
larva
Jan 11..
120
240
40
" 21
" 25 .
200
770
500
2,709
o
4,700
774
77
o
Feb 3
72
72
24
100
" 8
80
80
o
80
o
o
" 15 .
160
800
400
o
" 22. .
126
3,159
126
o
Mar 1
4
640
o
4
o
8. .
800
2 000
o
400
40
" IS. .
220
11 ,200
40
40
o
600
o
120
" 22
1 ,120
16,800
40
40
40
40
" 29
760
20 700
o
40
20
80
Apr. 5
" 12
1,420
1 100
32,300
25 100
20
60
o
40
40
40
80
160
" 19
" 26. .
5,500
24,320
27,200
20 800
o
o
100
320
100
300
300
May 3
19 600
182 400
o
o
o
o
400
" 10
53 200
169 600
o
o
200
400
" 17. .
27,900
166 400
o
900
4 800
200
700
" 24. .
12,160
126 400
o
840
40
80
280
" 31
5 680
103 800
o
160
40
80
440
June 7 . .
23,800
270 400
o
200
200
" 14
" 21
1 1 , 400
14 600
164,800
65 600
o
o
o
200
o
o
400
" 28. .
4 500
30 400
o
o
o
300
100
400
July S
" 12
1,320
780
2,400
180
o
60
o
40
180
480
300
" 19. .
1 ,680
10 800
o
60
o
80
480
" 26. .
840
11 600
o
o
o
120
Aug 2 .
360
11 200
o
80
o
40
80
9. .
400
2 400
o
o
" 16. .
600
1 600
o
60
60
60
" 23
3 360
11 200
o
o
60
60
180
o
" 30 .
5 520
13 600
o
o
120
Sept 6 . . .
240
8,000
o
o
40
160
80
" 13
4 320
16 000
60
o
o
120
180
o
" 20. .
19 000
o
60
120
60
60
" 27
Oct 4
2 600
96,000
29 000
o
200
o
o
300
400
80
200
40
" 11. .
3 920
27 500
o
80
o
40
40
80
" 18. .
5 580
17 500
40
40
o
40
40
120
" 25
2 400
22 000
120
o
o
120
60
o
Nov. 1 ...
8. .
300
960
8,000
14,000
o
120
180
120
" 15
400
12 000
100
o
100
100
" 22. .
160
10 000
o
2 000
320
" 29. .
4
6 000
12
o
120
Dec 6. .
440
9 000
o
o
500
" 13. ...
1 ,060
30 300
60
o
" 20
1 800
20
o
o
o
o
" 27. .
920
5 400
o
20
40
4 780
36 707
11
10
39
318
76
124
(23)
340
TABLE I concluded.
ORGANISMS PER CUBIC METER IN PLANKTON OF ILLINOIS RIVER IN 1898.
(An asterisk at head of column indicates that all entries in it are based on
filter-paper collections.)
J898
li
o^J
H
Plumatella
statoblasts
Total
Glochidia
Total
Miscellaneous
Total
Phytoplank-
ton
Total
Zooplankton
!
^
c
cS rt
oE
h
Jan 11
6
120
200
300
200
40
120
40
60
160
60
180
40
400
80
800
40
40
1,640
80
900
220
400
320
100
1,200
100
60
80
60
80
60
40
120
80
500
20
40
154
24
80
160
40
40
100
20
200
300
120
120
400
60
100
240
180
60
80
80
1,220
7,720
7,738
648
2,160
6,000
3,285
1,124
3,360
3,280
1,600
2,020
1,860
980
3,100
20,160
10,800
13,600
213,900
5,360
4,720
24,000
4,400
3,300
10,900
5,800
5,320
2,680
1,440
3,440
840
6,580
8,420
5,320
- 4,320
2,420
5,920
12,100
2,740
1,240
1,140
3,860
4,360
15,300
7,500
25,680
1,900
680
1,560
320
340
544,201,080
202,136,180
180,295,247
619,030,830
297,341,760
653,122,000
115,514,812
21,790,800
96,590,840
118,382,400
127,930,360
53,463,040
42,470,860
90,934,320
927,956,220
3,872,537,280
3,200,166,960
4,467,165,760
2,148,960,400
190,671,160
252,704,250
259,129,000
1,149,333,480
641,056,900
612,686,240
257,668,840
223,936,060
1,578,635,720
281,604,120
369,169,240
3,084,000,880
583,940,376
544,041,260
797,312,600
488,146,040
676,773,060
330,837,120
511,099,300
264,225,400
126,398,510
182,891,160
218,768,810
209,418,675
277,953,180
407,573,600
466,411,780
364,032,900
529,250,270
1,715,442,415
848,243,820
387,414,000
1,042,720
288,340
195,484
1,381,960
651,200
1,091,920
715,697
1,809,688
542,680
453,100
484,760
444,900
363,980
1,432,400
2,134,800
16,092,840
898,919,800
42,826,200
31,091,900
4,969,580
2,772,200
9,695,000
10,959,400
15,159,600
7,796,700
495,337,320
4,135,160
1,717,240
907,240
5,833,440
69,874,760
1,240,920
1,519,140
597,880
60,463,480
1,712,720
32,466,660
3,201,200
2,035.720
1,495,880
1,967,020
2,453,060
1,189,380
1,281,220
'732,300
1,109,040
799,980
916,780
1,757,440
682,440
548,700
545,243,800
202,424,520
180,490,731
620,412,790
279,992,960
654,213,920
116,230,509
23,600,488
97,133,520
118,835,500
128,451,120
53,907,940
42,834,834
102,366,720
930,091,020
3,848,630,120
4,099,086,760
4,509,991,960
2.180,052,300
195,640,740
255,476,450
268,824,000
1,160,292,880
656,216,500
620,482,940
753,000,160
228,071,220
1,580,352,960
282,511,360
375,002,680
3,153,875,640
585,181,296
545,560,400
797,910,480
548,609,520
678,485,780
363,303,780
514,300,500
266,261,120
127,894,390
184,858,180
221,221,870
210,608,055
279,234,400
408,305,900
467,520,820
364,832,880
530,167,050
1,717,199,855
848,926,260
387,926,700
" 21
" 25
Feb 3 . .
8
" 15 .
" 22. .
Mar 1
8. .
" 15
" 22
" 29 .
Apr 5
" 12
19. .
" 26. ...
May 3 . .
" 10. . .
" 17
" 24
" 31. .
" 14 .
" 21 ..
" 28
July- 5. .
" 12..
" 19
" 26
Aug 2 . . . .
9
" 16
" 23. .
" 30. .
Sept. 6
" 13. .
" 20
" 27
Oct 4. .
" 11
" 18
" 25. .
Nov 1
8. .
" 15. .
" 22
" 29..
Dec. 6
" 13
" 20. .
" 27
Average
37 135
52
9,393 723,283,871 34,226,468 756,548,801
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EXPLANATION OF PLATES.
PLATE I.
Seasonal distribution of synthetic groups of planktonts, Chlorophycece,
Bacillariacea, and Mastigophora, from July 1, 1895, to October 171896. Note
changes of scale indicated at bottom of diagram. Numbers in column at left apply
only to 1895. In this plate and in II. and IV., apices exceeding the limit of the
diagram are dropped down between dotted lines to show location. Circles at
bottom indicate location of day of full moon.
PLATE II.
The same as above, from July 1, 1897, to April 1, 1899. Note change in
scale from previous plate.
PLATE III.
Seasonal distribution of total Rotifera and Crustacea from July 1, 1895, to
October 1, 1896. The Crustacea included, belong almost exclusively to the Ento-
mostraca. Apices exceeding the limits of the diagram are dropped down between
dotted lines to show location. Totals include both adult and immature stages of
the Entomostraca when detached from parent, and both free and attached eggs of
the Rotifera.
PLATE IV.
The same as above, from July 1, 1897, to April 1, 1899.
PLATE V.
Seasonal distribution of Polyarthra platyptera. Total number of individuals,
not including eggs, represented by ordinants, parts of which exceeding 200,000
are represented by diagonal lines instead of solid vertical lines. Thus parts of a
seasonal plot which overlap those above it on the plate are represented by the
diagonally-lined ordinants.
(24) 355
356
357
358
359
360
ERRATA AND ADDENDA.
Page 58, line 7, for ovalis read ovata.
Page 85, line 8, for longicaudus read longicauda, and just above Phacus pleuro-
nectes read the following paragraph:
Phacus longicauda var. torta, n. var. This variety, for which I propose the
name torta because of the twisted body, is figured by Stein ( '78, Taf7207 Fig. 3). It
occurred sparingly in midsummer from July to September, rarely in October, in
.1896 and 1897.
Page 91, line 18, after T. caudata Ehrb. read T. lagenella Stein.
Pages]153, line 3 from bottom, 168, line 16, and 178, line 14, for '98 read '98a.
Pages 156, line 11, 159, line 16, and 161, line 5 from bottom, for '93 read '98a.
361
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