THE common: AED SUBSEQUENT GROWTH OF ‘

BAH-{MA PULEX AND D. MAGNA, POPULATIONS IN ’
FOUR INTERCOEéNECTED LAKES, . ‘ '

Thesis for the Degree of M. s. '
MICHIGAN STATE UNEVERSITY ‘
JAMES STEPHEN WEBBER
1974

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ABSTRACT

THE COLONIZATION AND SUBSEQUENT GROWTH OF
DAPHNIA PULEX AND Q. MAGNA POPULATIONS
IN FOUR INTERCONNECTED LAKES
By

James Stephen Webber

Four newly constructed interconnected lakes were filled
with water in autumn, 1973. A daily flow of treated sewage
was never achieved during the sampling period and the lakes
remained isolated from each other. Three lakes receiving
several tons of introduced aquatic macrophytes in October

were rapidly colonized by Daphnia pulex. Significant quanti-

 

ties of male 2. pulex appeared in November, apparently in

response to decreasing water temperatures. Population densities

declined following spring maxima. Reductions in most in-

stances were subsequent to reductions in pH or dissolved oxygen

concentrations. 2. magna appeared in Lakes 1 and 2 in July.
*ngheir increasing density and the coinciding density decrease

of 2. pulex might support the argument that Q. magna are more

efficient filter-feeders at higher temperatures.

of)
/\

£067

U

THE COLONIZATION AND SUBSEQUENT GROWTH OF

DAPHNIA PULEX AND 2. MAGNA POPULATIONS

 

IN FOUR INTERCONNECTED LAKES

By

James Stephen Webber

A THESIS

Submitted to
Michigan State University
in partial fulfillment of the requirements

for the degree of
MASTER OF SCIENCE
Department of Zoology

l9Th

ACKNOWLEDGEMENTS

I gratefully acknowledge the guidance of Dr. T. Wayne
Porter, chairman of my committee. His knowledge of
microcrustaceans, his much-needed criticism, his moral support,
and his patience over my two years of graduate work will
always be remembered with much gratitude.

I am very grateful also to Dr. Peter I. Tack and
Dr. Clarence D. McNabb, committee members, for their advice.
Their expertise in aquatic biology was greatly appreciated.

Special thanks are due the Michigan State University
Institute of Water Research for making the Water Quality
Management Project available for my research. I am especially
grateful to Mr. Joe Ervin for his assistance with equipment
on the site and to Mr. Charles Tanner for the analysis of.
chemical parameters.

A special thanks is also due Mr. Robert Glandon for
making available the data on the introduced aquatic macrophytes.

A final very special thanks goes to my wife, Nancy, whose

encouragement and support enabled me to achieve this goal.

ii

TABLE OF CONTENTS

Page

LIST OF TABLES . . . . . . . . . . . . . . . . . . . . v
LIST OF FIGURES. . . . . . . . . . . . . . . . . . . . ‘ vi
INTRODUCTION . . . . . . . . . . . . . . . . . . . . . 1
STUDY AREA . . . . . . . . . . . . . . . . . . . . . . h
METHODS. . . . . . . . . . . . . . . . . . . . . . . . 11
Physical and Chemical Monitoring . . . . . . . . . 11
Quantitative Site Sampling . . . . . . . . . . . . 11
RESULTS. . . . . . . . . . . . . . . . . . . . . . . . 18
Physical and Chemical Monitoring . . . . . . . . . 18
Temperature. . . . . . . . . . . . . . . . . . 18
Dissolved Oxygen . . . . . . . . . . . . . . . 18

pH 0 I O O O O O O O O O O O O O O O O O O O O 18

Soluble phosphorus . . . . . . . . . . . . . . 18
Inorganic carbon . . . . . . . . . . . . . . . 18
Ammonia. . . . . . . . . . . . . . . . . . . . 18
Quantitative Site Sampling . . . . . . . . . . . . 23
Lake

Lake

Lake
Lake

26

C’UJI'DH

DISCUSSION 0 O 0 O O O 0 0 0 O O 0 0 O O 0 O 0‘ O 0 O 0 31‘
SUMMARY AND CONCLUSIONS. . . . . . . . . . . . . . . . hi

BIBLIOGRAPHY . . . . . . . . . . . . . . . . . . . . . N3

iii

page

APPENDICIES. . . . . . . . . . . . . . . . . . . . . . hS
Appendix A: Chemical Parameters . . . . . . . . . AS
Appendix B: Daphnia spp. Population Parameters. . . 53

iv

Table
1.
Al.
A2.
A3.
Ah.
A5.
A6.
A7.
A8.
B1.
B2.
B3.
Bh.
B5.
B6.
B7.

B8.

Dissolved oxygen saturation

Chemical

Chemical

Chemical

Chemical

Chemical

Chemical

Chemical

Chemical

LIST OF TABLES

parameters:
parameters:
parameters:
parameters:
parameters:
parameters:
parameters:

parameters:

Lake

Lake

Lake

Lake

Lake

Lake

Lake

A

Top._.
Bottom
Top. .
Bottom
Top.

Bottom
Top. .

Bottom

Daphnia spp. population parameters:

Daphnia

Daphnia
Daphnia

spp.

spp.

population parameters:

population parameters:

pulex

 

Daphnia

pulex

 

Daphnia

pplex

 

Daphnia

pulex

 

Daphnia

pulex

 

population
population
population
pOpulation

population

parameters:
parameters:
parameters:
parameters:

parameters:

Site

Site

Site

Site

Site

Site

Site

A

B

Site C

Page
35
AS
A6
AT
h8
A9
50
51
52
53
5h
55
S6
57
58

59
6O

Figure

1. Water Quality Management Project .
2. Lake 1: Sites A and B

3. Lake 2: Sites C and D

N. Lake 3: Sites E and F

5. Lake A: Sites G and H

6. Daphnia pulex. . . . .

7. Daphnia magna. . . . .

8. Physical and chemical parameters:
9. Physical and chemical parameters:
10. Physical and chemical parameters:
11. Physical and chemical parameters:
12. Daphnia spp. population parameters:
13. Daphnia spp. population parameters:
1h. Daphnia spp. population parameters:
15. Daphnia spp. population parameters:
16. Daphnia spp. population parameters:
17. Daphnia spp. population parameters:
18. Daphnia spp. population parameters:
19. Daphnia spp. population parameters:

LIST OF FIGURES

 

 

vi

Lake 1.
Lake 2.
Lake 3.
Lake h.
Site
Site
Site
Site
Site
Site
Site

Site

Page

13
15
19
2O
21
22
2h
25
27
28
29
3O
32

33

INTRODUCTION

This investigation was designed to determine the response
of selected freshwater filter-feeding planktonic micro-
crustacean species to nutrient levels in the water. These
nutrients are first assimilated by phytoplankton or bacteria
and these organisms thus become a food form utilizable by
filter—feeding zooplankton.

Two major groups of freshwater planktonic microcrustaceans
are filter—feeders: calanoid copepods and cladocerans.
Calanoid copepods are generally found in the limnetic habitat
of larger lakes. Consequently they were not expected to
appear in abundance in the small lakes investigated. Clado—
cerans are common in both limnetic and littoral habitat.

This investigator studied the ubiquitous limnetic cladoceran,
Daphnia spp.

Zooplankton feeding has been reviewed extensively
(Conover, l96h: Edmondson, 1957: Jorgensen, 1966). Many
investigators have considered the role of cladocerans as
herbivores, grazing primarily on phytoplankton. Saunders
(1969) contended that thin-walled phytoflagellates are most
digestible compared with heavy—walled algae and blue—green
algae which may pass through the cladoceran intestine and

remain viable. Ingestion, assimilation, survivorship, and

reproduction of D. pulex fed blue-green algae were lower than

those fed green algae (Arnold, 1971). Furthermore, some bluee

green algae exhibited toxicity or inhibition toward D. pulex.
Recent investigations have indicated that cladocerans

are not dependent solely on algae for nutrition. ITaub and

Dollar (1968) found Chlorella Sp. and Chlamydomonas Sp. to

 

be deficient in meeting the nutritional requirements of

Q, pulgx. While detritus has normally been considered non—
nutritious (Conover, l96h), Pennak (1955) contended that
detritus must be the most important cladoceran food source

in the Colorado lakes he studied. Artificially created
detritus, labelled with carbon—1h, comprised half of the
organic matter assimilated by Daphnia spp. in Frains Lake,
Michigan (Saunders, 1969). Bell (1970) estimated that contri—
bution of detrital carbon to daily incorporation of Daphnia
pulex was much less than contribution of phytoplankton but
was significant. Incorporation of phytoplankton was more
efficient than that of artificial detritus, as was also shown
by Saunders (1969).

Bacteria have also been considered important in cladoceran
diet (Manuilova, 1958; Rodina, 1958). Bacteria seemed to be
limiting to zooplankton production in eutrophic lakes investi-
gated by Hilbricht g£_al. (1966). Sorokin (1957) reported
a correlation between the distribution of Daphnia spp. and
chemosynthetic methane oxidizing bacteria in a reservoir.

A general correlation between zooplankter size and bacterial

assimilation capacity was noted by Saunders (1969). Monakov
and Sorokin (1961) contended that zooplankton are more
efficient in assimilating algae than bacteria.

In any case, diet of cladocerans is complex.. While
phytoplankton may be the major component in the diet, detritus
and bacteria, while more refractory to digestion, can be
important supplementary components. They may even be dominant
in eutrophic situations where there are high concentrations
of bacteria and detritus.

This investigation was originally designed to correlate
ig_§i£u feeding rates of cladocerans with their population
densities and growth rates in each lake. Algal and bacterial
species found in the lakes were to be carbon-1h labeled and
used in a grazing chamber to detect feeding behavior differ-
ences between lakes. But continuous waterflow never occurred,
the lakes remained isolated, and a nutrient gradient never
developed. Hence the feeding experiment was not executed.

In mid—July, l97h, a leak was discovered in one of the
artificial marshes adjacent to Lake 3. Lake 3 was drained
and in early August Lakes 1 and 2 were also drained. The
leak was repaired but the ponds were not completely filled
again until early September. Cladoceran populations in the
newly refilled lakes would likely have been influenced by
variables not present in the lakes in July. Consequently

sampling was terminated in late July.

STUDY AREA

The study area is located at the intersection of Jolly
and College Roads on south campus of Michigan State University
(Figure 1). Construction of four interconnected lakes was
directed by the Institute of Water Research of Michigan
State University.

The lakes were designed to receive two million gallons
of secondarily treated sewage per day. Each lake is situated
at an elevation lower than the preceding lake and gravity
creates waterflow between lakes. Water flowing from Lake 2
can be passed directly through a water main to Lake 3 or be
diverted into or through three artificial marsh areas west
of Lake 3. Water from Lake A may be pumped to experimental
Spray irrigation sites. Water levels, flow rates, and
avenues of flow are directed and determined in a control
house on the east shore of Lake A.

Surface areas of the four lakes vary from 8.1 to 12.3
acres. The lakes are shallow, with maximum depths of 2.1
to 2.h meters limited to narrow channels draining into the
outlets (Figures 2—5). The lake bottoms are sealed with clay
to prevent seepage and consequent contamination of the ad«
Jacent ground waters.

Water was initially passed into Lake 2 in early October

of 1973. Upon its filling, four aquatic plant species were
A

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collected from four sites across Michigan and were introduced
for management purposes. Lakes 3 and h were the next lakes
to be filled with water and were also planted with macro-
phytes. Lake 1 was the last lake to receive water and no

aquatic plants were introduced.

METHODS

PHYSICAL AND CHEMICAL MONITORING

Thirty—one different physical and chemical parameters
'were monitored monthly by the Water Quality Laboratory of
Michigan State University. Determinations were made from
water samples taken near the surface and bottom of each lake.
Parameters given in graph form in this investigation are the

mean values of the top and bottom measurements.

QUANTITATIVE SITE SAMPLING

Sampling sites were selected that appeared to be best in
detecting responses to nutrient gradients within the lakes
system. Each lake was assigned two sampling sites. Sites
were chosen at a depth of 1.8 meters near the inlet and out-
let of each lake (Figures 2—5). Duplicate samples, which
should be the minimum allowable for population studies, were
taken at each station from depths of O, 0.6, 1.2 and 1.8
meters.

Zooplankton were trapped in a 2.1 liter horizontal
transparent van Dorn water sampler. Utilization of a towed
sampler, such as a Clarke-Bumpus plankton sampler, while
superior in quantifying zooplankton densities, would have
been impractical in the shallow lakes among the dense aquatic

macrophytes.

ll

12

Samples were collected in a three hour period at midday.
Since cladocerans tend to clump near bottom during daylight,
the integrating process will tend to smooth out these aggrega-
tions (Hrabacek, 1966). Furthermore, vertical distribution of
cladocerans was determined to be of no significance to this
research. Hence, samples from the four depths were integrated
into one sample.

In early May, larger, stronger-swimming cladocerans were
observed swimming to the bottom of the sampler.- By remaining
below the spigot, they avoided inclusion in the sample.‘ Sub-
sequently, the bottom trap of the sampler was removed and its
entire contents poured into a plankton tow net. The samples
from the other three depths were likewise poured into the net
and samples thus integrated in the field. Previously, samples
from the four depths were poured into separate containers and
integrated later in the laboratory. All samples were preserved
in 95% ethyl alcohol for later quantification. Identifications
were made according to Ward and Whipple (1959).

All Daphnia spp. in each integrated sample were counted.
Then approximately 100 were measured for length and their eggs
counted. Because 2, pulex (Figure 6) and D. magna (Figure 7)
are non-cyclomorphic cladocerans, length was measured from
anterior-most point of helmet to base of caudal spine, and was
determined to the nearest 0.1 mm. 97 percent of Q. pulex with
broods were 1.6 mm. or longer. Thus 2. pulex less than 1.6 mm.
in length were classified as Juveniles. Broods were observed
only in Q. magna larger than 2.6 mm. Consequently, D. magna

2.5 mm. or less in length were classified as Juveniles.

13

Figure 6. Daphnia pulex. A) Gravid female, lateral view, x38.

 

B) Male, lateral view, x6h.

C) Postabdomen, female, x80.

 

15

Figure 7. Daphnia magna. A) Gravid female, lateral view, x28.

B) Postabdomen, female, x53.

 

 

 

l6

 

17

Edmondson and Winberg (1971) have reported success in
maintaining eggs and embryos within cladoceran brood pouches
by dipping the living organisms in 95% ethyl alcohol. Be-
cause this method was not totally successful for this
investigator, all eggs in the subsample were counted, whether
they were inside or outside the maternal brood pouch. Fully
developed 2. EELSE young larger than 0.5 mm. in length were
observed in brood pouches. Consequently, all 2, pglg£
smaller than 0.5 mm. in length were considered prematurely
ejected embryos and were counted as eggs. Fully developed
2. magna were not seen in brood pouches.

Reproductive rates were calculated using Edmondson's

(1968) egg-ratio equation:

.E;
B = D
where:
B = average brood size
E = eggs/adult female
D = development time of eggs in days

Finite birth rate (b) can be calculated from B according
to Hall (196A):

b = ln(1+B)

Wet weights of Daphnia spp. were determined according to
Burns' (1969) equation:

w = 0.0116 Lb 2-67

where:

W = wet weight in mg.

Lb = body length (excluding caudal spine) in mm.

RESULTS

PHYSICAL AND CHEMICAL MONITORING (Figures 8—11)

Temperatures rose slowly in all four lakes from January
to March. Upon the disappearance of ice cover, temperatures
increased sharply until June at which time they began to
level off at approximately 20°C.

Dissolved oxygen concentrations in Lakes 3 and A declined
gradually from peak values under ice until May and June when
decline became more pronounced. Lake 1 and 2 were charac—
terized by more dynamic fluctuations in dissolved oxygen
concentrations.

In Lakes 1, 3, and E pH increased slowly to greater than
9.0 but dropped sharply to as low as 7.A in Lake A in
April and in Lakes 1 and 3 in May. Lake 2 was characterized
by a sharp pH increase in January following a slow decline,
but pH dropped suddenly in April and continued declining.

Soluble phosphorous concentrations in Lakes 1, 2, and 3
rose under the ice cover until January when concentrations
began a steady decline. Lowest concentrations occurred in
May, after which concentrations began to increase. Lake A
was characterized by a very gradual decline throughout
sampling period, although a minimum of 0.13 mg/l—P did occur

in May.

18

l9

 

 

 

 

‘I 3.0
'00
IO #3
' SOIUIlI
monoamc .0
PHOS'HOIUS
CAGION ‘0 ( I”
3'
(mull ‘ "o
10
‘7 I l l l l l l I l 0
0C! NOV BIC JAN "I MA. A'I MAY JUN “ll
3. " 7 "i 2.
24 F‘ ' d 14
20 P '4 20

VIA‘PIIAYUIE 1gp- “‘6 DISSOlVID

'C

 

 

 

 

 

 

 

 

I? d" OXYOQN
(MOIH
4 44
o 1 1 1 0
OC! NOV DIC JAN "I MAI APR MAY JUN JUl
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9L AMMONIA
'" - l o
:1 ' (ms/I)
. .. : — —. _
fl
7 .4
J 1 J l J l l l J OJ

OCT NOV DEC JAN FEB MAI APR MAY JUN JUl

Figure 8. Physical and chemical parameters: Lake 1.

20

 

- 3.0
I00
.0" ’
00 . , a A 2.0
monomc “‘ SOlUIll
‘° _ mosmonus

. -I.0 (MOI!)

CAIUON
40
("'0") /

 

 

I I I I I I I I I

 

 

 

 

 

 

0
OC' NOV BIC JAN III MAI AII MAY JUN JUI.
2. - -
1 2.
2‘ I- .-I( 2‘
20 P - 20
YIMPIIAYUII 1‘ I- "I I‘ DISSOIVED
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C I! I- .4 II OXYOIN
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I I I I I I I I I ‘ OI
ocr NOV 0!: JAN "I MAI API MAY JUN JUl

Figure 9. Physical and chemical parameters: Lake 2.

 

 

 

21

 

 

 

 

 

 

 

 

 

 

 

 

1 3.0
I00.
00 .
, sown!
INOIOANIC ‘° mosmouus
“no" 40 m (Moll)
(Moll) ‘ ° _. _ ..
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ac: NOV 0!: JAN "I MAI An MAY JUN JUL
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I
20 ' CI 2.
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00! NOV 00: JAN "I MAI API MAY JUN JUI.
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‘ I J L L I l I I O"
0:! NOV 00: JAN In MAI An MAY JUN JUL
Figure 10. Physical and chemical parameters: Lake 3.

 

 

22

 

 

'1 3.0
100 -
00 I- «A 2.0
sown!
INOIOANIC 50 - PHOSPI-IOIUS
CAIION
40A- 41.0 Ins/I)
(mall) \ __ _ _
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0 I I I I I I I "1 "" I 0
ocr NOV 0!: JAN "I MAI API MAV JUN

 

 

 

 

 

 

 

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24 I- .124
20— - 20
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(ma/I)
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4 - 4
0. I I I
on NOV DIC JAN "I MAI APR MAY JUN
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fl 9 - AMMONIA
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7 +- \ I
J b d
I I I I I I I I J 0"
oc: NOV 0!: JAN n0 MAI API MAY JUN
Figure 11. Physical and chemical parameters: Lake A.

23

Total inorganic carbon concentrations in all four lakes
fluctuated throughout the sampling period but minima for
each lake occured in March.

Ammonia concentrations decreased in all four lakes
during the sampling period. The decrease was gradual except
for a Sharp decline which occurred in all four lakes in

April.

QUANTITATIVE SITE SAMPLING

Mean lengths and variances of the duplicate samples were
compared to determine any Significant differences between
the duplicate samples. Employing student's t-test with a
.01 level of significance, determination was made that there
was insufficient evidence to indicate a difference in 85
percent of the duplicate samples.

Daphnia spp. were not detected in Lake 1 at either site
in autumn, 1973 (Figures 12 & 13). Dense pOpulationS
( 106/m3) of rotifers in the genera Asplanchna, Brachionus,
and Platyias in April were succeeded by a rapid population

growth of Daphnia pulex by the first week of May. From this

 

peak, the cladoceran populations slowly declined. In the
second week of June, all 2. pulex from Lake 1 were pink.

2. magna first appeared in samples in the first week of July,
comprising eight percent of cladoceran population. By the
last week in July, 2. magna comprised 96 percent of Daphnia

population.

war
was"!‘ ‘

(mlm’)

 

ID

I I TITI" I l I TUTTI! I 1 T1 III“ I I] I III"

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1° 7—44
OCT NOV DEC JAN

TOO

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Juvsmuss
so
(ab)

25

 

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105

I l IIIIII

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II IIIIII

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Figure 12.

OCT "NOV DEC JAN FEB MAI API MAY JUN JUI.

Daphnia spp. population parameters:

.0001

Site A.

 

 

 

25

    

 

 

 

 

 

 

IO = 105
E I“ r~
L. I I
,IO‘: . : '0‘
E. Z
r- -4
WET .. .
WEIGH! ‘ DENSITY
”3’: 5”: (m4)
(mo/m3) E II _....
- --1
r -
I02: 310’
E 2
p -1
IO‘h—l—J—k II I I J I ‘0‘
0C! NOV DEC JAN FEI MAI APR MAY JUN JUL
IOOP -1
75- .1
JUVENILES I
50- .01
(0M (Cass/9)
25?- ".001
I I I I I I I I I .0001
OCT NOV DEC JAN FEB MAR APR MAY JUN JUt
Figure 13. Daphnia spp. population parameters: Site B.

26

Fluctuations in B values generally preceeded correspond-
ing fluctuations in population densities. Percentage of
Juveniles in populations were generally inversely proportional
to B values. Only one male 2. pulex was observed in Lake 1
during sampling period and no male 2, magna were seen.

2. pulex population densities measured at sites C and D
in Lake 2 increased into the second week in November and
subsequently declined (Figures 1h & 15). Densities increased
in spring and a maximum was reached in the first week of
May. Following a decline, a second peak of same magnitude
occured in the first week of July. E. magna appeared by the
last week of July and comprised one percent of Daphnia
population.

Spring and summer values of B at site C generally de-
clined and had little correlation with population densities.
Fluctuations of spring and summer B values at site D approxiw
mated fluctuations of population density.

Male 2. pulex comprised from 8.3 to 25 percent of the
population in autumn but were not detected the following
spring or summer. No Q. magna males were observed. Peru
centage of Juveniles in the population was generally
inversely proportional to B values.

2. pulex populations in Lake 3 at sites E and F reached
peaks in the second week of November (Figures 16 & 17).
Population at site E achieved two density maxima in May and
June. Population at site F reached a single peak, of
lhO,OOO/m3, highest density for any site, in the third week

of June. 2. magna were not observed in Lake 3.

 

 

 

 

 

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‘OILU_L I I I I I I I I ‘0‘
OCT NOV DEC JAN "I MAI APII MAY JUN JUI.
I00!- I
75*- .I
JUVENIlES .
50L .0!
I°I°I I ' (099“?)
25,, I .001
I
I
_.I._ I I J I I .0001
OCT NOV DEC JAN FEB MAR APR MAY JUN JUL
Figure 1h. W spp. population parameters: Site C.

28

     
 

 

 

 

 

 

 

no’: :105
: , '1
I I ’ \ :
- I‘ I \“
\ I Q
- I I
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ocr NOV DEC JAN FEI MAI API MAY JUN JUI.
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.000I
OCT NOV DEC JAN FEB MAI API MAY JUN JUL

Figure 15. Daphnia spp. population parameters: Site D.

 

 

29

 

 

 

 

 

 

 

I05: 3 I05
I I
I I
b d
b a
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OCT NOV DEC JAN MAI A'I MAY JUN JUL
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(0].) \_l (“99”?)
25b fiDOI
/
/
. — I I _I L I I I J .0001
OCT NOV DEC JAN FED MAI A'I MAY JUN JUI.
Figure 16. Daphnia spp. population parameters: Site E.

 

29

 

 

 

 

 

 

 

 

ID _-_- 3 105
; :.
I- 1
I0”: : ‘0‘
I 1
b J
p-
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2
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OCT NOV DEC JAN FEB MAI API MAY JUN JUI
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(ole) \_l ('SQ’IQI
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, —‘ I I J I I I I I .0001
OCT NOV DEC JAN FEB MAI API MAY JUN JUI.
Figure 16. Qgphgia spp. population parameters: Site E.

M

 

 

30

 

 

 

 

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I I
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d
: "1
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OCT NOV DEC JAN FEB MAI API MAY JUN JUL
I00 - I
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$0
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(0M (Coos/9)
25 ‘IDOI
I I I I

 

OCT NOV DEC JAN FEB MAI API MAY JUN JUL

 

 

 

 

.000I

Figure 17. Daphnia spp. population parameters: -Site F.

31

At both sites B values varied generally inversely to
densities.. Fluctuations of Juvenile percentage at_site E
preceded fluctuations in densities. Fluctuations of
Juvenile percentage at F did not correlate with fluctuations
of population parameters at the site.

2. EELSE males were present in fall populations in Lake 3,
comprising l0.5 to 16.7 percent of the population, but only
one male was detected the following spring and summer.

2. pulex populations in Lake h at sites G and H increased
in densities during the third week of November (Figures 18 &
l9). Densities increased again in April and subsequently
declined sharply during the first week of May. Densities
fluctuated to two more maxima at site G and rose sharply to
one maximum at site H. 2. magna were not detected at either
site.

At site G, B values and Juvenile percentage fluctuated
concurrently but did not correlate with density. B values
and Juvenile percentage at site H were not so closely cor-
related and neither one correlated closely to fluctuations
in density.

2. pulex males comprised up to 33 percent of the popula—
tion at site H in November and were present at both sites
early next spring but had disappeared by the third week of

May.

32

  

 

 

 

 

 

 

 

 

 

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50- .01
(0M (0993/?)
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I J l I I I I I I 000'

OCY NOV DEC JAN FEB MAR APR MAY JUN JUL

Figure 18. Daphnia spp. population parameters: Site G.

M

 

 

.33

   
 

 

 

 

 

 

 

 

IO : 105
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I y’ -
'04: I .10‘
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25*- \ “.00I
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Figure 19. Daphnia spp. population parameters: Site H.

DISCUSSION

Sudden reduction in ammonia and subsequent declines in
pH seen in all four lakes in spring could be associated with
primary productivity. Peak primary productivity in early
spring, characteristic of most freshwater bodies, creates
a high pH. High pH will convert much of the inorganic
nitrogen to ammonia, much of which is subsequently lost to
the atmosphere as a gas. As a result of this large inorganic
nitrogen loss in the four lakes, primary productivity may
have become nitrogen—limited. Decreased photosynthesis
results in decreased pH while oxygen production is drastically
reduced.

Soluble phosphorous and inorganic carbon concentrations
never fell below .13 mg/l—P and 15 mg/l-C and probably
neither were limiting to photosynthesis.

Dissolved oxygen concentrations less than 8.0 mg/l
cannot be ascribed to lowered solubility due to increased
temperatures in these lakes. In temperature ranges en—
countered, dissolved oxygen saturation values exceed 8.0 mg/l
at an elevation of 900 feet (Table 1). Oxygen is being
depleted by some type of demand, probably respiratory.

In Lake 1, the lowest dissolved oxygen concentration for

any of the four lakes, 3.8 mg/l, is recorded in the third

3h

I" Il‘ s...» .‘ H u: "afiwfimfl

Table l.

35

Dissolved oxygen saturation in water at 900 ft.

elevation.

-< ——_—~fi4—_7
__#__—_-m

36

 

 

Table l.
TEMPERATURE 100% SATURATION
(0C) mg/l - D.0.
15 9.5
16 9.3
17 9.1
18 8.9
19 8.8
20 8.6
21 8.5
22 8.3
week of June. The concentration may have fallen even lower
between May and June monitoring dates. In this case,

appearance of pink-colored 2, pulex at sites A and B in the
first week of June coincides with low dissolved oxygen con»
centration. Some pond cladocerans produce increased amounts
of erythrocruorin when dissolved oxygen concentrations are
low (Pennak, 1953) , giving the organisms a pinkish color.
Apparently low dissolved oxygen concentration in Lake 1
initiated such a response in Q. pulex.

Oxygen reduction may have caused the concurrent D.
pulex population crash at site B. A similar crash occurred
at site A but not until a month later. Two factors may have
contributed to the difference in time between the two sites.
Site A, situated in a more open area of Lake 1, may have
experienced more mixing by wind and subsequently slower

decline in oxygen. Site B is adJacent to the outlet channel

lllll'il‘l‘IIIlII

 

37

of Lake 1. The lake bottom slopes toward this relatively
deep channel and gravity may have caused particulate organics
to aggregate in the bottom of the channel. Site B, then,
could have experienced greater oxygen loss due to increased
biological oxygen demand in the nearby channel.

None of the other lakes experienced so severe an oxygen
depletion nor the appearance of pink 2. pulex.

At five other sites (C, D, F, G, and H), declines in D.
23155 densities are preceded by lowering pH. Decreases in
pH may be indicative of reduction in primary productivity
and consequent decline in algal populations. Assuming algae
to be a maJor component in the diet of Q. pulex in these
lakes, g. p313; densities would decline as their food
disappeared. Such may be the case at these five sites.
Likewise in Lake 1 this phytoplankton reduction may have
interacted with low dissolved oxygen concentrations to cause
the crashes in the D. pulex populations. Declination of
population density at site E preceded pH declination.

Predation probably was not an important facter in re-
duction of cladoceran densities. Cyclopoid copepods,
ubiquitous predators on rotifers and planktonic micro—
crustaceans, were never abundant in samples taken from the
lakes. Larval Chaoborus spp., predators common in limnetic
habitat, were detected in only a few samples. Failure to
detect these two invertebrate predators in quantity, however,
may have been due in part to sampling techniques. The

horizontal sampler employed in this investigation creates

38

turbulence as it is lowered through water and cyclopoid

copepods and larval Chaoborus spp., being faster swimmers

 

than Daphnia spp., may have escaped capture.
Fish, particularly small ones, are important plank-
tivores. Some fish species were probably introduced with

the introduction of macrophytes in October, 1973. Mosquito-

 

fish, Gambusia affinis, were extremely abundant in the small
original marsh 100 feet south of Lake 3 and could easily have
been carried from the marsh to any one of the lakes on the
feet and feathers of waterfowl. However, offspring of a

few pioneer fish would not have been able to fill the lakes
with significant populations in less than a year. Hence the
impact of fish on the cladoceran populations is considered

to be slight.

Fungal and/or other diseases may have contributed to
high density losses observed in populations, but cladocerans
were not examined for diseases.

Juvenile percentage B, and population density should
have been more closely correlated within sites. According
to Wright (1965) increased brood sizes would have predicated
increased densities, a correlation detected clearly only at
sites A and E. Hall (196h) reported that a high relatively
constant proportion of Juveniles is reflective of a popula-
tion growing under relatively stable conditions. Such a
high relatively constant proportion of Juveniles was present
at sites C, D, and E but in these cases population densities

l

were fluctuating.

39

Colonization of the lakes by 2. pulex was rapid. One
possible source of this species was the original marsh.
Plankton samples were taken from the marsh in June, 1973,
when construction of the lakes was in its initial stages.
The only planktonic microcrustacean was a single copepod
nauplius. 'Abundance of planktivorous mosquitofish in the
marsh undoubtedly kept water depleted of most large zoo-
plankton. Mud samples from the marsh were cultured in
laboratory and Q. pulex were hatched, probably from ephippia
observed earlier in mud samples.

A more probable source of large numbers of Q. pulex
was the several tons of aquatic plants introduced to the
lakes in October. Subsamples of the plants were cultured in
laboratory and Q. pulex were produced in subsamples from
three of the sites. The potentially large number of Q.
pulex included in several tons of wet macrophytes could
easily account for the rapid colonization of Lakes 2, 3, and
h in the autumn. Lake 1 was not planted with macrophytes
and D. pulex did not appear there until the next spring.

Appearance of 2. magna in Lakes 1 and 2 in July is
interesting from the view point of their filtering rates.
Burns (1969) determined that adult 2. pulex filtering rate
was slightly higher at 20°C than at either 15 or 25°C. In
contrast, filtering rate of 2. magna increased with increasing
temperature and at 25°C was more than twice its rate at

15°C. This increased filtering capacity of D. magna at

hO

higher temperatures would increase its capability of dis-
placing 2. pulex in summer.

In Ward and Whipple (1959) the appearance of males in
normally parthenogenetic female populations is reported to
be initiated by a shortage of food. In Pennak (1953)
production of males is attributed to other factors:

1) crowding of females and a subsequent accumulation of
waste products, and 2) decreasing water temperature.

2. pulex populations in the lakes had reached only low
densities in autumn when significant numbers of males appeared.
Consequently there could have been little metabolic waste
product accumulation in the lakes. Food was probably plenti—
ful to the sparse populations, especially inthe form of
suspended organic detritus which would have abounded in the
newly—filled lakes. Falling water temperature, then, would

best explain male production.

SUMMARY AND CONCLUSIONS

While the scope of this investigation was less extensive
than originally planned due to complications beyond the
control of this investigator, there were someresults of
interest: 1) Colonization of three of the lakes by D. pulex
was rapid. The apparent sources of D. pulex were the intro—
duced macrophytes. 2) D. pulex began to disappear as
D. magna populations grew during the summer. 3) Predation
was apparently not important in reducing cladoceran popula—
tions during the sampling period. h) Appearance of D. pulex
males in autumn was apparently initiated by falling water
temperatures. 6) None of the four lakes were identical in
fluctuations of physical and biological parameters.

This investigation pr0posed to correlate fluctuations in
some population parameters of Daphnia spp. with changes in
physical and chemical parameters. The lack of well-defined
correlations between these parameters indicates the im-
plausibility of correlating zooplankton response directly
to physical and chemical variations. An investigator cannot
concentrate on a single parameter or a narrow set of para-
meters to the exclusion of others. All parameters of both
biological and physical nature must be considered in natural

population studies.

kl

A2

The four lakes were similar in many respects. They
shared the same geographical location, they were constructed
at the same time, they were similar morphologically, and
three were filled in the same month with the same type of
water and were planted with aquatic macrophytes. In spite
of these similarities, the lakes varied greatly in both
physical and biological characteristics. This variation
would reinforce the argument that results of limnological
research in one situation should be extrapolated only with

greatest caution to other situations.

BIBLIOGRAPHY

BIBLIOGRAPHY

ARNOLD, D. E. 1971. Survival and reproduction of Da hnia
pulex fed blue green algae. Limnol. Oceanogr. 16:906-920.

BELL, R. K. and F. J. WARD. 1970. Incorporation of organic
.carbon by Daphnia pulex. Limnol. Oceanogr. 15:713-726.

 

 

BURNS, C. W. 1969. Relation between filtering rate, tempera-
ture and body size in four species of Daphnia. Limnol.
ggganogr. lhz693-700.

CONOVER, R. J. 196A. Food relations and nutrition of zoo-
plankton. Symp. on Exp. Mar. Ecology, Univ. Rhode
Island, Occas. Pub. 2:81—91.

 

EDMONDSON, W. T. 1957. Trophic relations of the zooplankton.
Trans. Amer. Microscop. Soc. 76:225-2h5.

 

 

EDMONDSON, W. T. 1968. A graphical model for evaluation the
use of egg ratio for measuring birth and death rates.
Oecologia 1:1—37.

 

EDMONDSON, W. T. and G. G. WINBERG. 1971. A manual on
methods for the assessment of secondary productivity in
fresh waters. Blackwell Scientific Publications, Oxford
and Edinburgh, Great Britain. 358 pp.

HALL, D. J. 196h. An experimental approach to the dynamics
of a natural population of Daphnia galeata mendotae.
Ecology h5:9h-112.

HILLBRICHT~ILKOWSKA, A., Z. GLIWICZ, AND J. SPODNIEWSKA. 1966.
Zooplankton production and trophic dependencies in the
pelagic zone of two Masurian lakes. Verh. Int. Verein.
Limnol. 16zh32-hh0.

HRBACEK, J. 1966. A morphometrical study of some backwaters
and fishponds in relation to representative plankton
samples, in Hydrobiological Studies, Hrbacek, J. (ed.),
pp. 221~256.

JORGENSEN, C. B. 1966. Biology g£_Suspension Feeding.
Pergamon, N. Y. 313 pp.

h3

hh

MANUILOVA, E. R. 1958. The question of the role of bacterial
numbers in the development of cladocera in natural con-
ditions. Dokl. Biol. Sci. 120:h38-hh1.

MONAKOV, A. v. and Y. I. SOROKIN. 1961. 'Quantitative data
on the feeding of Daphnia. Trud. Inst. Biol. Vodokhr.
h:251-261.

 

.PENNAK, R. W. 1953. Fresh-Water Invertebrates of the United
States. Ronald Press Co., New York. 769 pp.

 

PENNAK, R. W. 1955. Comparative limnology of eight Colorado
mountain lakes. Univ. Colorado Stud., Ser. Biol.
2:1—75.

RODINA, A. G. 1958. Microorganisms and increase of fish
production in ponds. A. 1, Moscow. 171 pp.

SAUNDERS, G. 1969. Some aspects of feeding in zooplankton.
In Eutrophication: Causes, Consequences, Correctives.
Nat. Acad. Sci. Publ. 1700. Washington, D.C.

 

 

SOROKIN, Y. I. 1957. The role of chemosynthesis in the
production of organic matter in water bodies. Mikrobiology

26:736-7hh.

 

TAUB, F. B. and A. M. DOLLAR. 1968. The nutritional inadea
quacy of Chlorella and Chlamydomonas as food for Daphnia
pulex. Limnol. Oceanogr. 13: 607 —611.

 

WARD, H. B. and O. C. WHIPPLE. 1959. Fresh-Water Biology,
Second Edition, (W. T. Edmondson, ed.7 John Wiley and
Sons, New York, N. Y. 12h8 pp.

WRIGHT, J. C. 1965. The population dynamics and production
of Daphnia in Canyon Ferry Reservoir, Montana. Limnol.
Oceanogr. 10:583—590.

APPENDICES

APPENDIX A

CHEMICAL PARAMETERS

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30 3.0 0.0.0 .000 mu fig - hand-u.

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odé 0.? 0.0x 0.“ 0.) uo - 333353

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row 4..» .00 .0». .00 <§ .hv fl? «2 (as - 53.68
3.0 0.0 .+\ $ .0 1M\ .r .3 x (as - 53.53qu
44.0 mmd mad #0 34 an; t.« 4.3...“ a (as - 0538 -magocamo;
find 30 no.0 find ‘3; nfix hug“ 3 a (we - P30730232“.
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no.0v no.0 n0.0v $0.0 3.0V 3.0V to QUE . Eat—Haw...

.mm H» .sfim .mm iwmr .02 .m3 5 Que - onto—5
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u H\me - uwcmmgo um>Pomm_o-coaLmo

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m0.0v n.0.0v «0.0V 00.00. 00.0... 00.0v 3.0V 3 :03 n 53503

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.5 .2 .3 .T .0 .0. .0 0 “NW5 . 0.2030 .307000000

.01 .0. u :05 .. 0200.5 0030350023

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0.02 0.02 0.02 0.0 0... oo - 0.230.202.2223

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2. 0.0 20.0 No.0 .nl0..0 00¢ «0.0 .30 0&0 a (as . 0333 -miofimoi
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02.0w .02 .000 .000 000 .02m .0002 .0002 Euhsco .. 00:30:02.8
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0 .\ms - uwcmmgo 00>.omm_o-congmu

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Mud m6 Nud add $20 3.0 Iwmd 3.0 2.0 3; a <9: .. pmuopumagofimof
dnwww mwdw. mmavmv .mmVMV .un.m. o<.o .mmwmw mmuu. him. :

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APPENDIX B
DAPHNIA SPP.

POPULATION PARAMETERS

53

 

acmns.qm mom .xmpam.qm.u¢.«.
2%... .la. a 33% 4m as:

xm_=m.qm meagfi

 

 

 

 

m.mm “Ho. 0 ww.o mo.H_ mm.owm.e .¢.mmH.NH «**¢~ Paw RN
Em mmo. o mm.o om.H o~.~¢~.m .«moo.- .«¢N _=a No
oe mHo. eo.o N~.o _ em.H oo.o¢a.m *mom.~m .eu and «H
_mm owe. o Rm.o on.” oe.~mo.m «mum.nm «¢~ saw He

m.me NmN. o mm.o mo.H mm.m-.oH *mm¢.m~ «es Au: HN
“a Nmo. o Nm.o mm.o m~.H~N *ooo.e «cm A”: co
. a *eu La< HH

o *mk >02 on
o Auo._v$ mxadv *mN >02 mg
----- mu poo mm
----- m“ “co m”
----- mu goo HH
any any cowumw>mo Ams\mev
mm_w=m>=q m mm_mz ugaucmum cgmcoA com: .92 um: me\*

< muwm "mgmumsmgma cowumpaaon .anm mwcnmmo .Hm anmp

su

 

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20000.4. 0000 0

Aumcpmgv mxmdv

 

 

 

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0.00 000. 0 00. 00.0 00.000.00 00000.00. 00 000 00
0.00‘ 000. 0 00. 00.0 00.000.0 0000.0 00 000 00

00 000. 0 00. . 00.0 00.000.0 0000.00 00 000 00
0000 000. 0 00. 00.0 00.000.0 0000.00 0> 202 00

00 ~00. 0 00. 00.0 00.000 0000.0 00 202 00

. 0 00 000 00

0 00 >02 00
0 00 >02 00
2000000 02000
----- 00 000 00
----- 00 000 00
----- 0 00 000 00
200 200 000000>00 200\000
000000>00 0 00002 00000000 000000 0002 .02 002 m0\0

m 000m ”0000050000 00000—30oa..qqm 0000mmo .Nm mpnmh

55

 

 

 

 

 

, M-
040E430 .flad000 00
.mmflmm.qm 0000 0
00000000 00000
00 0 0 00. 00.0 00.000.0 00000.00 00 000 00
0.00 . 000. 0 00. 00.0 00.000.0 0000.00 00 000 00
0.00 000. 0 00. 00.0 00.000.0 0000.00 , 00 000 00
00 000. 0 00. 00.0 00.000.0 0000.0 00 000 00
00 000. 0 00. 00.0 00.000.0 0000.00 00 202 00
00 000. 0 00. 00.0 00.000.0 0000.00 00 202 00
.0 000.0 0 00. 00.0 00.000 0000 00 000 00
00 000. 00 00. 00.0 00.000 0000.0 00 >02 00
00 0 0.00 00. 00.0 00.000 0000.0 00 >02 00
000 0 00 00. 00.0 00.00 0000.0 00 000 00
000 0 0 00. 00.0 . 00.0 0000 00 000 00
0 0 0 0 0 0 0 00 000 00
000 000 000000>00 0000000
000000>0w m 0000: 0000:00m 000000 :00: .03 003 me\0

.0 000m 00000050000 0000000000 .000 00:;m0o .mm 00000

1.. 1.1.3.3.. .
ll... .. .. . 50%

56

 

Avo000su 02000
0

 

 

 

 

00 0 _0 00. 00. 00.000 000.00 000 00
0.00 000. 0 00. 00.0 00.000.0 000.00 00 000 00
00 000. 0 00. 00.0 00.000.0 000.00 00 000 00
00 000. 0 00. 00.0 00.000 000.0 00 020 00
00 000. 0 00. 00.0 00.000.0 000.0 00 202 00
00 000. 0 00. 00.0 00.000.00 000.00 00 202 00
2 . 000.0 0 00. 00.0 00.000 000 00 000 00
00 000. 0.00 00. 00.0 00.000 . 000.0 00 >02 00
00 0 0.00 00. 00.0 00.000 000.0 00 >02 00
00 000. 0.0 00.0 00.0 00.000.0 000.0 00 000 00
0 0 0 0 0 0 . 0 00 000 00
0 0 0. , 0 0 0 0 . . 00 000 00
any ARV 000000200 Ams\msv
000000>20 0 00002 00000000 200000 0002 .02 002 0000

.0 000m 00000050000 0000000000 .000 00::m00 .00 00000

57

 

----- A 00 000 00
00000000 02000

 

 

 

 

0.00 000. 0 00. 00. 00.000.0 000.00 00 000 00
00 000. 0 00. 00.0 00.000.0 000.00 00 000 00
0.00 000. 0 00. 00.0 00 000.0 000.00 00 000 00
0.00 000. 00. 00. 00.0 00.000.0 000.00 00 202 00
000 000. 0 00. 00.0 00.000.00 000.000 00 202 00
00 000. 0 00. 00. 00.000 000.0 00 000 00
00 .000. 0.00 00. 00. 00.000 000.0 00 >02 00
00 0 0.00 00. 00.0 00.000 000.0 00 >02 00
00 0 0.00 00. 00. 00.00 000.0 00 000 00
000 0 0 00. 00. 00 00 000 00 000 00
0000000 02000

----- 00 000 00

000 000 000000>00 0000000

000000>20 0 00002 00000000 000000 0002 .02 002 0000

 

.0 000m 00000050000 0000000000 x0030 0000000 .00 00000

58

 

 

m.0w men. 0 we.
m.nm moo. 0 mm.
m.mm o 0 mm.
m.hm .NNO. 0 mm.
mm one.. 0 mm.
NW cm“. 0 em.
0 o 0 HH.
mm o m.mH mm.
mm Ham. 0 wH.H
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0m—_:w>:w m mapmz vgmucmum

 

00.0
vo.0
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mm.~
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mm.
m~.H
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cameo; com:
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HH.mm~.m
oo.~nm.m
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mn.mm¢
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cm.~00
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oom.mu
mHm.me

wme.o¢~
moo.on

mum.mn
mmm.¢
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mm“

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em
0
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000 00
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000 00
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.mm mpnmh

59

 

 

 

 

 

0.00 000. 0 00. 00.0 00.000.0 000.00 00 000 00
00 000. 0 00. 00.0 00.000.0 000.00 00 000 00
00 000. 0 00. 00.0 00.000.0 000.00 00 000 00

0.00 000. 0 00. 00. 00.000.0 000.00 00 000 00
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0 0 0 0 0 0 0 00 >02 00
. 0000000 02000
---- 00 000 00
---- 00 000 00
---- 00 000 00‘
000 000 000000200 000\000
000000000 0 00002 00000000 000000 0002 .02 002 00\0

 

.u mp0m ”wgmpmsugoa 2000002000 x0020 0022000 .00 0000»

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NN. 0m. . mm.mmm.H mwm.w~ . 00 03w 0N

 

 

00 000. 0
00 . 000. 0 00. 00.0 00.000.0 000.00 00 000 00
00 000. 0 00. 00.0 00.000.0 000.00 00 000 00
0.00 000. 0 00. 00. 00.000.0 000.00 00 000 00
0.00 000. 0 00. 00.0 00.000.0 000.00 00 202 00
00 020. 0.0 00. 00.0 00.000.0 020.00 00 202 00
00 000. 0.0 02. 00.0 00.000 000.0 00 000 00
000 0 00 00. 00. 00.00 000 00 >02 00
0000 000. 00 00. 00.0 00.02 000 00 >02 00
0000000 00000
----- 00 000 00
----- ‘ 00 000 00
----- 00 000 00
000 000 000000200 000\000
000000200 0 00002 00000000 000000 0002 .02 002 00\0

 

.: 000m "0000050000 2000002000 2002a 0022000 .wm 00000

HICHIGRN STATE UNIV. LIBRARIES

 

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