FOREST SUCCESSION ON‘ THE
POORLY-DRAINED SOILS IN THE HIGGINS
LAKE AREA OF MICHIGAN

Thai: for flu Degree of Ph. D.
MICHIGAN STATE COLLEGE
Tap Ku
I954

This is to certify that the
thesis entitled

”CREST SUCCESSION 0N POORLY DBAIm SOILS

IN HIGGIRS LAKE AREA 01' MICHIGAN"
presented by
TAO EU

has been accepted towards fulfillment
of the requirements for

__2h.D_._ degree in M

 

____.—-‘——-—

 

 

AUG I475: 01
y 03

 

OVERDUE FINES:
25¢ per day per Item

RETURNING LIBRARY MATERIALS:

Place In book retum to remove
charge from c1rcu1at1on records

 

Ill ‘llfl'll' l'lllll‘i'lvlll,

IX '1

FOREST SUCCESSION ON THE POORLX-DRAINED SOILS

IN THE HIGGINS LAKE AREA OF MICHIGAN

I"
’5

I
A
1

BY
TAO*KU

.-

A THESIS

Submitted to the School of Graduate Studies of Michigan
State College of Agriculture and Applied Science
in partial fulfillment of the requirements
for the degree of

DOCTOR OF PHILOSOPHY

Department of Forestry

1954

y. .

l“ n

o

s.

.11: n‘noa F

..‘e‘ “abfio a: r:
R
i

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fl:

Tao Ku
Candidate for the degree of

Doctor of PhilOSOphy

Firual examination, Friday, July 30, 1:30 P.M., Forestry
Building

Dissaertation: Forest Succession on the Poorly-drained Soils
in the Higgins Lake Area of Michigan.

Outline of Studies:

Major subject: Forestry
Minor subjects: Soils, Geology

B iographic a1 Items :

Born, March 26, 1925, Chaochow, Kwangtung, China

High School, Nankai Middle School, Chungking,
Szechuan, China, 1943

Undergraduate Studies, University of Nanking,
Nanking, China, 1943-1948-B.S. degree,Forestry

Graduate Studies, Michigan State College,
1949-1950 - M.F. degree, Forestry
1950-1954 - Michigan State College,

East Lansing, Michigan

Experience: Member, Chinese Youth Expeditional
Army (203D), 1945

Member of Xi Sigma Pi

343484

The anti:
itpreciation to I
Farestry, uniep ‘

I 1.1 . ,
3113‘s; tars i}

n. “.8 val‘lab‘£
:clr. Egbert E
"”39 Of he .
u,

IVntlcted.

ACKNOWLEDGMENTS

The author wishes to exPress his sincere thanks and
appreciation to Dr. T. D. Stevens, Head of the Department of
Forestry, under whose inspiration, supervision, guidance and
orifizicism this investigation was undertaken and completed.

The writer is further indebted to Dr. William B. Drew,
Dr. lKirk Lawton, and Dr. Stanard G. Bergquist for their kind
guidance, helpful suggestions and criticism in the conduct of
this work. .

Grateful acknowledgment is also due to Dr. C.L. Gilly,
for his valuable help in identifying the vegetation specimens;
UDIDr. Robert E. Dils, for his criticism in the manuscript;
and for the assistance received fromer. H. V. Borgerson, in
dharge cf the Higgins Lake State Forest where this study was
conducted.

The investigator extends his sincere thanks to
Mr. Kim K. Ching, who is working on Forest Succession on the
Upland Soils in the same area, for his helpful assistance and
BuSgestions in the field, and the continuous discussion there-
after in conducting this investigation.

To my wife, Victoria Ku, who contributed to certain
PhaBes of the manuscript, and her continuous encouragement

and Support, grateful acknowledgment is given for her patient
effOrt.

FOREST 5';
IN THE

‘
V“.
~‘ .LI‘

Dyed

FOREST SUCCESSION ON THE POORLX—DRAIMED SOILS
IN THE HIGGINS LAKE AREA OF MICHIGAN

By
TAO KU

IAN ABSTRACT
Submitted to the School of Graduate Studies of Michigan

State College of Agriculture and Applied Science
in partial fulfillment of the requirement
for the degree of
DOCTOR OF PHILOSOPHY

Department of Forestry

Year 1954

Approved /{ he leéfiéfi/I

' a
testifv the vari

iraized soils in
;::table trend of

The H13?
Ital part of the
.. is within the
21:: as defined

F

Soils 1
TV

~¢3W3ap COTij
‘§« I
kaI‘azs 0O “ ‘
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“H, 1/”; acre.
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it

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V

The objective of this study is to distinguish and
identify the various types of vegetation found on the poorly—
drained soils in the Higgins Lake area and to determine their
probable trend of succession upon the ecological basis.

The Higgins Lake area is located in the northern cen—
tral part of the lower penisula of Michigan. Ecologically,
it is within the Hemlock-White Pine-Northern Hardwoods Forma—
tion as defined by Nichols (1935), Braun (1950), among others.

Five types of vegetation were recorded on the poorly-
drained soils in the area. They were: (l)The Marsh or Open-
meadow type, (2)the Swamp Shrubs of Sglig:00rnus-Alnus Associ—
ation, (3)the Lowland Aspen of Populus-Salix Association, (4)
the Swamp Coniferous forest, and (5)the Swamp Hardwood forest.
Quadrats of three sizes were used for the vegetation study,
1.9-, 1/5 acre quadrat for tree vegetation; 1/10 acre quadrat
for Shrub vegetation; and l/lOOO or milacre quadrat for vege-
tation below six feet tall, the herbaceous layer.

Ecological factors were considered in three groups:
(1)311matic, (2)Edaphic, and (3)Biotic. Among the climatic
factors discussed, evaporation is believed to show the com-
tfined effect of the other three, temperature, relative humi-

dity, and precipitation. Data of evaporation rates in three

 

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vi
TAO KU ABSTRACT
different vegetation types were obtained by using Livingston
atmometers. Results show that the evaporation rate decreases
fronxthe Marsh to the Swamp Shrubs type and to the Swamp Forest.
This indicates that although the forest cover may be the cause
of the lower evaporation, the higher evaporation rate in the
Marsh may still be an important factor in retarding the estab-
lishment of forest cover on that area.

Soils Of all the quadrats were investigated. Samples
of seven soils were collected by horizon to determine their
Characteristics. Statistical analyses were applied to analyze
all the laboratory data. Results show that there were no dif-
ferences between the volume weight and the total porosity of
the soils. Only the Newton sand had a lower cappilary poro-
Bity than the others. Results of field determinations of soil
mollture leads to the belief that soil moisture content decre-
ases from the Marsh to the Swamp Shrubs and to the Swamp Coni-
rePous Forest. The peaty soils had higher organic contents
than the mineral soils. Soils of the Marsh type were strongly
‘Old in reaction, followed by the two Rifle peat soils of the
Swamp Shrubs and the Swamp Conifers. The Bergland soils of
the Swamp Hardwood type were almost neutral in reaction.

Biotic factors are not believed to be of great signi-
ficance to influence the forest succession on the poorly-
whmined soils in the region. Fire which is usually caused by
man is probably the most important factor that will induce

secondary succes s ion .

uflain this a:
efsmzession is
Slants to Swamp
arelativel; lo:
:Iiax type of 1
labiland Asm
33%:

[porary t

to!
V1
'4

375-“ 0? otherwi:

vii

TAO KU ABSTRACT

Diagrams of forest succession on the poorly-drained
soils in this area is shown in Fig. 12. The principal trend
of succession is believed to be that from Marsh to the Swamp
Shrubs to Swamp Conifers to Swamp Hardwoods and finally after
a relatively long period of time to the upland, mesOphytic,
climax type of the Hemlock-White Pine-Northern Hardwoods.
The Lowland Aspen type is most commonly thought to be the fire
or temporary type after the original forest has been burned
over or otherwise denuded. The Swamp Hardwoods, the Swamp
Conifers of Thgjgrépigg-giggg, and the Marsh type are all

believed to be the physiographic subclimax type to the area.

TABLE OF CONTENTS

LISTFOF'TABLES ........................................
LIST OF FIGURES .......................................
LISTIOF PLATES ........................................
INTRODUCTION ..........................................
REVIEW OF LITERATURE ..................................
A.ECOLOGICAL CONCEPTS ................................

B.CLIMAX FOREST OF THE NORTHERN LOWER MICHIGAN .......

C.PERTINENT LITERATURE TO THE STUDY OF VEGETATION ON
POORLY'DRAINED SOILS eeeeeooeeeoeoeocoo-0000000...o

DESCRIPTION OF THE REGION eeeoeeeooooooooooeoooeoeoooeo
A0HISTORY AND GENERAL DESCRIPTION 00000000000000.0000.

B’sLIMATE oeeSeeoo0000.00.00.00.coco-09000000000000...
C

.GEOLOGY OOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOOCO00......

D

‘SOIL 000000000000000000000000000000000000000000.0000

742.com TYPE
VEGETATION ANALYSIS ...................................
A.FIELD PROCEDURE ....................................
B-OFFICE PROCEDURE ...................................
C-VEGETATION INVENTORY ...............................

1.8WAMP HARDWOODS OF BLACK ASH-AMERICAN ELM-RED
MPI—E ASSOCIATION 00.0.90...OOOOOOOOOOOOOOCOOOCO.

2.3WAMP CONIFERS OF BLACK SPRUCE-BALSAM FIR-NORTHERN
WHITE CEDAR ASSOCIATION 00.0.00...00.00.00.000...

BOLOWLAND ASPEN OF POPULUS’SALIX.ASSOCIATION 0000000
4.3WAMP SHRUBS OF SALIX-CORNUS-ALNUS ASSOCIATION ...

SQMARSH 00.0.0.0...0.0...COOOOOO0.000000000000000000

Page

xiii

XV

ID

10

15
24
24
30
34

2+3
47
2+7
51

56

68
80
88

 

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1x

Page

ECOLOGICAL FACTORS 99
IL.CLIMATIC FACTORS .................................. 99
_1.TEMPERATURE ..................................... 1OO
2.RELATIVE HUMIDITY ............................... 103
3.PRECIPITATION ................................... 104
4.EVAPORATION ..................................... 106
B.EDAPRIC FACTORS ................................... 113
1.SOIL PROFILE DESCRIPTION ........................ 114
2.SOIL VOLUME'WEIGHT .............................. 118
3.SOIL POROSITY ................................... 121
4.SOIL.MOISTURE ................................... 127
5.SOIL TEMPERATURE ................................ 132
6.SOIL ORGANIC MATTER CONTENT ..................... 133
7.SOIL REACTION ................................... 138
O.SIOTIC FACTORS .................................... 143
1.1NSECTS AND DISEASES ............................ 143
2.ANIMAIS.......................................... 146
3.MEN ............................................. 147
FOREST SUCCESSION .................................... 149

sIINICULTURAL CONSIDERATIONS ......................... 162
SUMMARY

00.0.0.0...OOOOOOOOOOOOOOOO....000.00.000.000. 165

APP‘j-‘TI-N’DIX 00.00.000.000...ooooooooooooooooo00.00.00.000 169
BIBLIOGRAPHY o00.00000000000000000000000.000.000.000... 179

PIATE 0......0.000.....00.00....COOOOOOOOOOOOOOOOOOO... 190

. . O -

p v
Nut

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Weather I

Sumzarize
Stands 1

Sumzary 1
0f the :
centaqe

'1'!
9.88899

Summary

by 3129'

Snmnary
0f the

SUEIapy
0f the
tags of
3138895

Sumzary
by SIZe

sumlarv
by 591;

Sullary
Of the
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NqueP

SUmlar
@1an S
Secorie
°tems

Suziary ‘
Pf the ;
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or 516:;

Table

10

ll

12

LIST OF TABLES

Weather Data of the Higgins Lake Area ...........

Summarized Data of Four l/5 Acre Swamp Hardwood
Stands by Size Classes .........................

Summary Data of the Vegetation in Milacre Quadrats
of the Swamp Hardwood Stands as Recorded in Per-
centage of Coverage or Number of Stems by Height

Classes 0.0.0.0....OOOOOOOOOOOOOOOOO0.00.0000...

Summary Data of Four 1/5 Acre Swamp Conifers Stands

by 8126 Classes O...0.0.0.0...00.000000000000000

Summary Data of l/lO Acre Quadrats by Height Classes

of the Swamp Conifers Stands ...................

Summary Data of the Vegetation in Milacre Quadrats

of the Swamp Conifer Stands as Recorded in Percen—

tage of Coverage or Number of Stems by Height

Classes 00......0.0.0.0...OOOOOOOOOCOOOOOOOOOOOO

Summary Data of Two 1/5 Acre Lowland ASpen Satnds
by 3126 Classes .OCCOCCOOOOCOOCCCOOCOOOCOQOOCOOO

Summary Data of TIO l/lO Acre Lowland Aspen Stands
by Height Classes 000000000000000000000000.0000.

Summary Data of the Vegetation in Milacre Quadrats
of the Lowland Aspen Stands by Height Classes.
Plants Vere Recorded in Percentage of Coverage or
Number Of Stems 0000000000.0.0000000000000000...

Summary Data of the Four Milacre Quadrats in the

Swamp Shrubs Stand by Height Classes. Plants Were

Recorded in Percentage of Coverage or Number of

Stems 0.00....OOOOOOOOOOOOOOOOOOOOOOOOOCOOCOOOOO

Summary Data of the Vegetation in Milacre Quadrats

of the Three Marsh Stands by Height Classes. Plants

Were Recorded in Percentage of Coverage or Number

Of Stems 000000000000.0000000000000000...00.0000

Weekly Averages of Temperature, Relative Humidity
and Accumulated Precipitation During the Period
from July 20 to October 4, 1952. Compiled from
Data Obtained at Weather Bureau Stations near the
Higgins Lake Area 000.000.0000..0.00.00.00.00...

Page

32

63

64

75

76

77

83

. 83

89

96

101

.........

......

.....

......

mayorati
of Three
Period f

Analysis
from Th:
July 2C

Volume 'a':
Soils

Result 0

Volume

PEPCenta
Horizon

‘n31YSis
and T01

Pore 3?.
80115‘]

Distri

3011 KO
f9 Pen+

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Table
13

14

15

16

17

18

19

2O

21

22

23

24

25

26

27

28

xi
Page

Evaporation Rate as Recorded by Week at Stations
of Three Different Vegetation Layers During the
Period from July 20 to October 4, 1952.......... 106

Analysis of Variance of the Evaporation Data Obtained
from Three Different Stands During the Period from
July 20 to October 4, 1952 ..................... 108

Volume Height of Each of the Three Horizons of Six

80118 .0...0..00...0.....00.0..0.00.0.00.000...0 120

Result of the Statistical Analysis of the Soil
Volume Weight Data 00.000.000.00...coco-0.00000. 120

Percentages of Soil Porosity of Each of the Three
Horizons Of 51X 80118 00.000000000000000000.0000 122

Analysis of Variances for Non-Capillary, Capillary
and Total Porosity of Six Soils ................ 123

Pore Space Distribution in Percentages of the Six
80118 by Horizon 0.000....0.0.....0...000..00... 125

Analysis of Variance from Data of the Pore Space
Distribution of the Six Soils by Horizon ....... 126

Soil Moisture Data of Rifle Peat Soils in Two Dif-
ferent Stand Types, Obtained During the Period from
July 20130 OCtOberz", 1952 .000000..0.0.0.00.00. 128

Analysis of Variance of the Moisture Contents of
the Two Rifle Peat 50118 0....00.0....0...0..000 130

Soil Temperatures of the Two Rifle Peat Soils Which
Were Measured Once Every Week During the Period
from Ju1y 20 to OCtOber 4, 1952 0000000000000... 132

Organic Matter in Percent of Seven Soils by Horizon
as Determined from Dry Combustion Method ....... 135

Analysis of Variance from the Data of Organic Con-
tent of Seven Soils by Horizon ................. 136

Soil Reaction (pH Values) of the Seven Soils by
Horizon .C....................0.000.....0000..00 139

Analysis of Variance for the Soil Reaction Data of
Five Soils by Horiaon .......................... 140

Composite Summation of Edaphic Characteristics of
seven 80118 by Horizon .0.00.0........0...0....0 142

‘‘‘‘‘‘

xii
Table Page
29 Data of Relative Frequency, Relative Density,

and Relative Basal Area of the Dominant Compo-
nents of the Swamp Hardwoods ................... 157

13m

1 X5? 0? lov
Khulna]
While F11
Relation

9 3‘18:- or s:
Cary N11
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Weather
lake Ar
Study ,

)5? of i
Surface

5 1"1519 or
SSPVln

Veek1y
Recori
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t0 03f

"eek1y
from.

weekly
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weekly
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Ferl<
8011 1

LIST OF FIGURES
Figure Page

1. Map of Lower Michigan Showing the Location of the
Higgins Lake Area in Relationship to the Hemlock-
White Pine-Northern Hardwoods region and also Its
Relation to Some of the Tree Ranges and Soils .. 25

22 Map of Higgins Lake State Forest Showing Its Boun-
dary Within the Four Adjoining Counties:Missaukee,
Kalkaska, Crawford and Roscommon................ 26

:5 Map of Lower Michigan Showing the Location of U. S.
Weather Bureau Stations in and near the Higgins
Lake Area Whose Weather Data Are Used in This

StUdy 000000000ooooeoooeoooeeoooeoooooeoeooooooo 33

‘1 ‘Map of the Higgins Lake State Forest Showing It8
surface GeOIOgy 0.00.00.............000.......0. 36

5 Map of Lower Michigan Showing Drainage Basins
serVIHS the Higgins Lake Area cocoon-0000000000. 37

5 weekly Data of Maximum and Minimum Temperatures as
Recorded at the Higgins Lake and Houghton Lake
Weather Stations During the Period from July 20
to OCtObeP 4, 1952 00000000000000.00000000000000 102

7 Weekly Accumulated Precipitation During the Period
from July 20 to October 4, 1952 ................ 105

8 Weekly Evaporation in Cubic Centimeters in Open-
Meadow, Swamp Shrubs, and Swamp Conifers Stands
for the Period from July 20 to October 4, 1952 . 107

9 ‘Weekly Evaporation (Average of Three Stands) in
Relation to Temperature and Precipitation for the
Period from July 20 to October 4, 1952 ......... 112

10 £3011 Moisture Curves for the Two Rifle Peat Soils 129

11 Weekly Data of Soil Moisture Content in Percent
of the Two Rifle Peat Soils by Horizon ......... 131
12 ?Diagram Showing Successional Relationships exhibi-
ted Between the Associations of Poorly-Drained
Soils in the Higgins Lake Area ................. 150

"wise
it’vl

D Phytogre
Conifer

M Phytogrs
Herivo<

 

 

xiv
Figure Page

13 Phytographs of the Important Species of the Swamp
Coniferous Forest Type ........................ 156

11+ Phytographs of the Important Species of the Swamp
Hardwoods Type ................................ 159

 

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LIST OF PLATES

Plate

1 Set Up of the Livingston Atmometer in A Marsh Stand.

Inflorescences of Sgirpus cyperigus Can Be Seen on
the Upper Part of the Photograph. Bulb Was Appro-
ximately 18 Inches Above the Ground ............

2 Set Up of the Livingston Atmometer in A Swamp
Shrubs Stand. Bulb Was Approximately 18 Inches
Above the Ground ...0......0.0.0..0.0.0...0....0

3 A Sgirpus-Typha Stand of the Marsh T pe. The Domi-
nance of Cat-tails (Iypha 1§tifolia Is Indicated
by Their Numerous Flowering Stems in the Fore-
Ground. An ASpen (Pogulug tremuloides) Stand Is

 

Page

109

110

in the Back Ground. In Betfieen, Willows (Salix sp.)

Are Grown in Low Bushes to Give A Patchy Appea-

rance 0....0....0....0.0.........0...0..00..0.0.

4 A Salix-Calamagrostig Stand of the Marsh Type. An
Aspen (Popglus tremuloides) Stand Is In the Back

Ground 00000000000000000.000000000000000...000700

5 Soil Profile of the Newton Loamy Sand In A Salix-
Cglamagrostis Stand of the Marsh Type During the
Dry Season in October. The Water-table Was Down
to A Depth of Approximately 2 Feet. The Organic
Peat Layer Reached Down to About 1% Feet........

5 A Typical View of the Swamp Coniferous Forest ...

7 External Appearance of the Elm—Soft Maple Stand
of the Swamp Hardwoods Type ....................

8 Ground Vegetation of the Elm-Soft Maple Stand of
the Swamp Hardwoods Type Showing Seedlings of
Maple, Elm, and Ash. Carex intumescens Can Be
Seen In the Fore-Ground With the Inflorescences
Shown In the Center Of the Picture..............

9 Soil Profile of the Bergland Loam In the Ash-Red
Maple-Elm Stand of the Swamp Hardwoods Type.
Water-Table Was At Approximately 10 Inches .....

190

191

192
193

194

195

196

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“er"“y- boae 1 1

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32.193 801‘s 1

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INT RODUC‘T I ON

The objective of this study is to distinguish and
identifygthe various types of vegetation found on the poorly—
drained soils in the Higgins Lake area and to determine their
PPObable trend of succession upon the ecological basis.

This study was begun in 1952. At that time, this study
"as intended to be entitled: "Forest succession on the poorly-
drained soils in the Higgins Lake State Forest, Michigan."
However, for administrative reasons, the H ggins Lake State
Forest is not existing at the present time. It has been divided
into two parts which belong to two separate State Forests: The
AuSable State Forest and the Houghton Lake State Forest.

The area formerly of the Higgins Lake State Forest in Roscommon
C°unty and Missaukee County is now a part of the Houghton Lake
State Forest, whereas the area in Crawford and Kalkaska Coun-
ties is now a part of the AuSable State Forest.

Therefore "the Higgins Lake area" as appeared in the
ti'C-le is equivalent to that of the former Higgins Lake Forest
P981cm. It is for the same reason that some of the maps pro-
du°ed and statements made remain to be referring to the Higgins

Lake State Forest area.

ideological C
Before
“'3! 0f veget
iii :‘elimitati
Warmly
{fetter records

‘.':.at

 

REVIEW OF LITERATURE

A.Ecological Concepts
Before the establishment of Ecology as a science,the
study of vegetation was unchanging. It was a mere description
and delimitation of plant communities as absolutely static.
Warming (1909) made a great advance by gathering to-
gether records of vegetative change or succession. He stated

that

"In early times plant societies were looked upon
as stable groups which were in a state of quiescence,
complete in their development, and peacefully living
side by side. In reality no such relations exist in
the plant world. Everywhere and continuously there
is going on a struggle between plant societies; each
individual society constantly strives to invade the
field of others, and each small change in the living
conditions immediately produces shiftings and changes
in the mutual relations of these group.

This dynamic view leads to the aarly works of Cowles
(1899,1901, 1911), who conceived the modern concept of suc-

c‘Pvlaion. He stated that

"At the close of the vegetative cycle there is the
final vegetative aSpect varies with the climate, and
hence is called a climatic formation. -e- The ultimate
~stage of a region is mesbphytic. The various plant
societies pass in a series of successive types from
the original conditions to the mesophytic forest.’

q

Beginni
Sleients includ
:epts of once 3
isirgle climax
other permanent
Tiltion to 011
carted aub- or
itiah between I
5913! those on

“flied soils .

Nichols

3

Beginning at about the same time, the publications of
Clements included much that served to shape our present con-
cepts of succession. His "monoclimax" theory is that "only
a single climax exists in a given climatic region, although
other permanent communities determined by other factors in
addition to climate may occur there also; these would be
termed sub- or proclimaxes." He also attempted to distin-
Q‘Jish between primary and secondary succession, the former
being those on newly formed soils, and the latter these on
denuded soils.

Nichols (1923) has defined plant association as "a
SPOup or community of plants which occupy a common habitat,
“1d are essentially similar throughout its extent in physio-
Bnomy, ecological structure, and floristic composition."
this concept of "association" is used in this paper.by the -
anthor applying to the various associations encountered.
Nichols recognized the plant association as the fundamental
unit, of vegetation whereas it parallels to the'Climax“ or
”7°Pmation“ by Clements. NichOls described succession simply
‘5 “the replacement of one plant association by another: and
pI‘QGelded to classify successions into progressive and retro-
8x‘eesives in regard to "Trend”; whereas there is no such a
t‘hing as retrogressive succession in Clement' concept. In‘
relation to “Climax", Nichols' idea was that the nature of

cliliiax in any area is controlled by geographic conditions,

9 both the 1
magnized by l
' "a.
156 British :.
5329169 (1935)
At the
"histwas heli
h3‘48?» 29 to 9.
the teachings
”u
“me “Ion t
999%

. 33.1021. 51

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4

thus both the climatic climax and physiographic climax were
recognized by him. This is essentially the same as that of
the British "Polyclimax' position under the leadership of
Tansley (1935).

At the donference of Plant and Animal Communities,
which was held at Cold Spring Harbor, Long Island, New York,
August 29 to September 2, 1938, Conard (1939) brought together
the teachings of various schools which have had special in—
fluence upon the description and classification of'nature
Vegetation. Six schools were considered by him: (l)The Zflrich-
Montpellier School :IKerner (1863) sketched with permanent
Precision the major lineaments of central Eiropean vegetation.
The conspicuous feature was the stability of the magnificent
Plant mosaic of central Europe and Alps when left undisturbed,
and the concept of association was developed. It was defined
by its floristic composition as a unit actually found in
nature, and upon which all phytosociological study centered.
(2)1113 Scandinavian School: Influenced by its geographical
1°°Iltion where the vegetation is that of the marginal lands,
Ttwfi oft repeated emphases were developed: (a)The soil is
the product of vegetation, and is independent of the nature
°f the substratum; (b)the foundamental unit of vegetation is
the layer or synusia“ (Conard, 1939). (3)1’he Danish School:
R‘unkiaer (1934) divided the species of the world into ”life-
r“fins". The percentage of species in each.,.;;roup drawn from

“‘11 over the earth gave the "normal spectrum" for the earth

99 a whole. T2
cal treatment 1
i193?) quoted :
phytosociologi:
zegetation ni

the first to r.

:9th 5
due Sur‘r
‘330c1at
s Bear“

1,, is

 

 

5
as a whole. Thus, the statistical method of phytosociologi-
cal treatment was developed. (4)The Russian School: Conard
(1939) quoted as Sukatchew (l93#) summarized that "Russian
phytosociologists were primarily interested in the Steppe
vegetation and its relation to forest“. The Russians were
the first to recognize the relation of soils to vegetation
and climate (Glinka, 1914). (5)The Chicago School: The con-
cept of succession dominated the Chicago School under the
leadership of Cowles (1899, 1901). Its prime object of the
study of vegetation was an explanation of the causes and
Processes of vegetational change. (6)The Nebraska School:
The concept of succession was deve10ped under the inspira-
tion of Clements, with its extensive terminology.

During the Conference, Gleason (1939) defended the
Individualistic Concept of the Plant Association. He stated
that the fundamental question basic to all ecological work
is, "what is a plant association?" In answering to this

A

question hepresented his theory:

”The vegetation unit is a temporary and fluc-
tuating phenomenon, dependent, in its origin, its
structure, and its disappearance, on the selective
action ofthe environment, and on the nature of‘
the surrounding vegetation. Under this view, the
association has no similarity to an organism and
is scarcely comparable to a species."

In defense this theory, the so called Individualistic

Concept, Gleason presented a series of theses, the main

paints were: (1)?
were in excess
93:9 nethod of '2
char station is
taticnal unit :39
theappearance 0
existing associa
531213 to clarif.
n .
F1?3b,3.‘
detached pie
community, ‘.

dimensions 1

119 its a:
theless 9. v;
Surveyed, :3}
area it cal:
uniformity :

 

 

poiths were: (1)Every species of plant has reproductive
Powers in excess of its need; (2)every species of plant has
seine method of migration; (3)the environment in any parti-
cular station is variable; and (4)the development of. a vege-
tational unit depends upon one or the other of two conditions,
the appearance of new ground, or the disappearance of the
existing association. Two general statements were introduced

by him to clarify these theses:

"First,an association, or better one of those
detached pieces of vegetation which we may call a
community, is a visible phenomenon. As such it has
dimensions and area, and consequently boundary,

Ihile its area may be large, the community is never-
theless a very tangible thing, which may be mapped,
surveyed, photographed, and analyzed. Over this

area it maintains a remarkable degree of structural
uniformity in its plant life. Homogenity of struc-
ture, over a considerable extent, terminated by
definite limits, are the three fundamental features
on which the community is based. Without these three
features, Grisebach would never have published his
statement a century ago; without them, all of our
studies of synecology would never have been developed.
Also besides its extent in space, every community has
a duration in time. Uniformity, area, boundary, and
duration are the essentials of a plant community.

Second, every community occupies a position in two
series of environmental variation. In the space
series, as the community exists here, in this spot,
it is part of a space-variation, and its environment
differs from the adjacent communities. In the time
series, as.the community exists pg! , at this time,
it is part of a time-variation and in its environment
differs from the communities which proceeded it or
follow it.“

~

The climax and its complexities have been ably dis-

cussed and summarized by Cain (1939). In his paper, he made

the following 8‘

'hlmlimax' t‘n.

"Since
Rietz, Ric
Scharfette
on the rec
be atresse
Climaxes 1

Clement
hF‘POthesis
different
rfixiizttions
chl‘onologi
disposed c
DOBtCIimay
pm381Ve I
climatic c

different
°" t0 dirt
Phillips.

(
“”1. 9st

things w}

 

 

the following statements concerning the "Monoclimax" and

"Polyclimax" theories :

"Since support of the polyclimax hypothesis (Du
Rietz, Nichols Tansley, Game, Gleason, Nordhagen,
Scharfetter, xen, Domin, Conard) rests largely
on the recognition of "edaphic climaxes", it should
be stressed that Clements too recognizes edaphic
climaxes in the concept of 'seration’.fl

Clements has met exponents of the.polyclimax
hypothesis in the following ways: (l)chorologically
different stable communities are considered to be
faciations and lociations of an association; (2)
chronologically different stable communities are
disposed of as subclimax, proclimax, preclimax,
postclimax, held in stability by a continuing re-
pressive factor; (3)stable communities not of the
climatic climax but due to biotic factors are dis-
climax, in part; (4)topographically different stable
communities, different because of continuing edaphic
factors are called serations; (5)variations due to
different soil types within a geological formation,
or to different geological formations, are recognized
(Phillips, 1934), apparently under the name of locia-
tion. The polyclimax view includes many different
things, while Clements uses a different term for a
different thing."

In his concluding statement, Cain pointed out the
difficulties in description and classification of vegetation
‘8 a result from lack of a standard terminology with the

following statements:

' ----- Clements' disposition of the variations

w1thin the climax (or climax region) through the
concepts of faciation and lociation, through sub-
climax, preclimax and serclimax, through aeration,
ItDd through preclimax and postclimax presents a-
8°ientifically and phylosophically sound system.
A.doscription and classification of all the stable
climax) communities of a region might necessitate
d°¢ling with all the above concepts. A very large
munberef investigators have chosen not to follow

1’" H O ('0 fl\;‘) [-9- O

a:

Clements' in this but to treat all such cover
types as 'associations‘ The plant sociologists
go even farther and include successional communi-
ties (associes) under the term. This may have
some Justification if the seral nature of the
communities is not proven. The result, however,
is to include many different things under the ‘
term, whereas Clements has a different term for
a different thing.“

‘

Curtis and McIntosh (1951) have prOposed the Concept
of"Vegetation Continuum for the upland forest of southwest

‘Wiaconsin. In their paper:

"The relative ecological importance of each
tree species in each stand was eXpressed by a
summation index of the relative frequency, rela-
tive density, and relative dominance herein
called the importance value.

Climax adaptation numbers were used to weigh
the importance value of each species in a stand.
The summation of these weighted numbers resulted
in an index which served to locate the stand
along a gradient. ----- the entire series of
communities formed a Continuum in which a defi-
nite gradient was exhibited from initial stages
composed of pioneer species to terminal stages
composed of climax species.”

Q

. .Braun (1935. 1950) has developed an interpretation
“ fill-Bax vegetation, the 'association-segregate' . It is a
climax'unit, which refers to space-time segregates of the
‘flncient mixed forest evolved by conditions of the environ-
“Wnt. and it includes consociation, faciation, and locia-
tion, regardless of the number of dominants in a stand.
1‘ h0P.Dociduous Forests of Eastern North America, Braun

has devoted a chapter to Forest Ecology and Terminology

because of the la
forest ecology .
Dansereau
tescription and r
basis. He sugges
tie: in floristic
51:91 classifioat

Wear shape , (5

 

 

because of the lack of uniformity in the terminology of
forest ecology.

Dansereau (1951) has proposed a new system for the
description and recording of vegetation upon a structural
basis. He suggested six criteria for the stddy of vegeta-
tion in floristic mapping, life—form statistics, and ecolo-
gical classifications: (l)life-form, (2)size, (3)function,
(4)1eaf shape, (5)1eaf texture, and (6)coverage.

ilhoax forest (

There ar-
heatate of £13?
sc‘itzem lower it
Firiuoodi Forest
:ezinsula and th

The 501“.

41561113 reoogni:

at 19 also 002120!

‘
shame

'9 °r this
than .

The fores

23:9 4‘
«0pm a Part

a

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’52::
rOI‘mgtion b h

 

lO

3.31imax forest of the northern lower Michigan

I There are two types of climax forest vegetation in
‘Une state of Michigan: the Deciduous rorest rormation in the
scuxthern lower renisuia, and the Hemlock-White Pine-Northern
Hardwoods Forest Formation in the northern part of the lower
peninsula». and 1~ throughout the upper. peninsula .

The boundary between these two formations is general-
ly"being recognized as near the latitude 43 degrees North.
It is also commonly described as a line running from Saginaw
to NMskegon. This boundary is not sharply defined and is re-
presented by an "ecotone" or a zone of tension. Baal and
Wheeler (1892), livingston (1903, 1905), Quick (1923),
Gleason (1924), Veatch (1952), Darlington (1945), Potder
(1946, 1948), Braun (1950), and others have pointed out the
Prosonce of this tension zone between the two forest formae
tions.

The forest vegetationwhuoh lies north of.thd’tension
zone forms a part.oftthe‘mixedfconifcntnonthern hardwood forest
formation. A number of descriptive names have been given to
thi' formation by many ecologists and botanists. The region
t° b0 studied in this paper lies within the "northern hard-
w°°d” r°egionolerothingham(1915), the northeastern "tran-
lition forest“ region of Nichols (1918), and the "Great lake"
<Ni"south Canadian forest" region of Hardy (1920). It is also

com"‘DOHded to the "lake forest“ region of Weaver d- Clements

{1929), the '89.“
the "St. Lawre a:
toe recently,
harlot (195C),
hoe-)iorthern E
Frothir
forest! as cos:
9! the norther:
between the no:
that the north.
Fraseoce of ye
‘53 the Ibsenc

"W31 other

".
m“ 1| more

11
(1929), the "eastern hemlock" region of Nichols (1935). and

the»"8t. lawrence-Great Lake” region of Harshberger (1911).
More recently, in Braun's Deciduous Forests of Eastern North
America (1950), the area is described as the Hemlock-White
Pine-Northern Hardwood region. ‘ I

Frothingham (1915) described the northern hardwood
forests as occupying "the fresh, well-drained, fertile soils
lof'the northern pine region". In discussing the differences
lmetween the northern and southern hardwood forests, he stated
'thst the northern hardwood forest is distinguished by the
presence of yellow birch, white pine and eastern hemlock,
and the absence of yellow peplar, red gum, sycamore and
several other more southern species. The northern hardwood
forest is more simple in composition and usually divided
into two regions: (l)the eastern mountain region, and (2)
the Great Lakes recion. He stated also that the greater
abundance of basswood and elm is perhaps the most striking
Characteristic of this forest formation in the Lake States.

Nichols (1935) designated this region as "eastern
hemlock-eastern white pine-northern hardwood region“ or
limply the "eastern hemlock region". He said that the climax
is a mesOphytic forest comprising a mixture of evergreen‘
coniferous and deciduous broadleaf trees. The characteristic
Species are: hemlock, sugar maple, beech, yellow birch, white

pine, basswood, American elm, white ash, red oak, black cherry,

12
red spruce, balsam fir, white spruce, red maple, and Norway
pine. He considered these trees into four groups with refer-
ence to their geographical distribution: (l)those whose
centers of north-south distribution are far to the south
(beech, white ash, black cherry, white elm); (2)those whose
centers are partly within and partly immediately to the
flouth (sugar maple, basswood,and northern red oak); (3)those
Whose centers are within the region and whose ranges are
more or less coextensive with it or some part of it (hemlock,
White pine, yellow birch, Norway pine, and red spruce); (4)
those whose centers of north-south distribution are to the
rlax-tin of the region (white and black spruce, fir, tamarack,
balssrn paplar, and paper birch).

Weaver and Clements (1929) described the "Lake Forest"
formation as consisting of ‘a single association in which white
pine, Norway pine and hemlock are the climax dominants.

A brief glacial history of the Lake Forest Formation
“0.3 presented by Potder (1946), in discussing the different
°p1nions to characterizing the climax of the "lake forest"
r‘99.;ien, he grouped the different interpretations into two
lines: "-----one favoring the pine-hemlock complex as climax,
the ether favoring hemlock-northern hardwoods as climax".
The latter gpo‘rpmhich includes Potzger, considered pines

Participating crown cover as relics, or post-climax.

...-..‘h-

lock-Yb?
fro: nor
through '
lantern

mthm

13

Braun (1950) has designated the extent of the Ben-
leck-White Pine-Northern Hardweed region as: "It extends
from northern Minneseta and extreme southeastern Manitoba
threugh the upper Great Lakes regien and eastward acress
southern Canada and New England, including, toward the
seutheast much of the Applachiam Plateau in New Kerk and
Northern Pennsylvania.” on. stated that the region is
characterised by the preneunced alternatiens ef deciduous,
conifereus, and mixed ferest communities. She listed ’
lugar maple, beech and basswood, sugar maple and beech, or
lugar maple std bassweed as:- the usual deminants, and yellow
birch, white elm and red maple mere or less frequent asso-
ciates in primary deciduous communities. The coniferous
cOmmunities which eocur at intervals almost throughout the
I‘Cgion are of two general types: (l)these of mere er less
dry sand plains and ridges where white pine, red, or Norway
Pine, and Jack pine prevail; and (2)these ef peerly-drained
‘I‘eas, begs er muskegs, where black spruce, northern white
°0dar, and tamarsok prevail..'rhe primary mixed communities
in this area are hemlock and northern hardwoods, principally
"agar maple, beech, bassweed, and yellew birch, in which
there is er was an admixture of white pine. These, the most
characteristic oemmunities give to the region its name,

Hemlock-White Pine-Northern Hardweeds.

B:
mis reg:
lathe Suj
Wreath:
“mm Uf
She state
1°“Steal c?
”31011 as
I»
SO'I'QVQP’
the We :
9. “
“equency
P361113 of
O
"r the p
1‘ fine-3
”521mm

“a Dem

14

Braun divided the Hemlock-White Pine-Norhhern Hard-
woods region into four sections: (l)the Great Lakes Section;
(2)the Superior Upland; (3)the Minnesota Section; and (4)the
Laurentian Upland Section. Northern lower Michigan and
eastern upper M1chigan fall into the Great lake Section.

She stated that beech-maple as a forest type or as an eco-
logical climax community is as well illustrated in this
region as it is in northern Ohio or southern Michigan.
"However, maple is generally more abundant than beech in
the more northern communities and usually has a higher
frequency.” She also pointed out that the sandy outwash
Plains of glacial topography afford suitable environment
for the pine forests and the morainal ridges if the soil
1! fined-grained and loamy are occupied by deciduous forest
°0lmunities or a mixture of deciduous species with hemlock
‘nd perhaps white pine. . '

Northern lower Michigan is a region of low relief
‘hd predominantly of glacial topography. There are extensive
‘Rndy eutwash plains and morainal ridges as well as innummer-
‘ble lakes, ponds, and poorly-drained areas. This paper,
‘3 the title indicates, deals with the forest succession on
Poorly-drained soils of area near the Higgins lake, therefore

1! limiting itsdlf to a much confined scope.

fl

3.?ertiner

drained

15
0.Pertinent literature to the study of vegetation on poorly
drained soils

Warming (1595) made variations in the water content

of the soil as a basis for clasSIfying.plant’societies;into
hydrOphytes, mesophytes, and xerophytes. Cowles (1901), after
efforts were made to classify the vegetation in northern
Michigan in 1898 in accord with Warming's principles, felt
the need of another classification.rAn.attempt was therefore
made by Cowles to relate the facts of distribution to com-
binations of factors and to group plant societies according
to their relationship and their evolution. He stated that
the climatic factors determine the distribution of plant
societies in large while local differences are produced by
changes in the edaphic or soil factors. Hence, he preposed

a l'physiographic classification” and stated:

"In a young tOpography, such as the recently
glaciated areas of Michigan, Wisconsin, and Min-
nesota, there is a great variety of tOpographic
conditions and of plant societies. Among these
are many hydrophytic lakes and swamps and many
xerophytic hills. In passing from youth to old
age then,,a region gradually loses its hydrOphytic
and xerophytic areas and the ultimate stage of a
region is mesophytic. The various plant societies
pass in a series of successive types frOm the ori-

ginal condition to the mesOphytic forest which may
be regarded as the climatic type."

q

Whitford (1901) followed Cowles' principle in his

study of the genetic development of the forests of northern

,A .

a
societies and 8‘
II: n
growth18 a ‘e'
Ecoloiic
northern lower 3
fists and ecologi
Gates (1925, 191
latte (1935-): 03
LiVlIlfiSt
tie: to be 618’“
Era'n'fori Countie
In a
\c}.uarack-l{or1
Eealso “‘
pointed

20a nearly the}

{tests due to t

6
t0 buhh

mlog near]

of distribution

16
Michigan. He pointed out the successive stages of the swamp
societies and stated that in each series the climax plant
growth is a "deciduous-hemlock combination".

Ecological studies on bog or swamp communities in
northern lower Michigan have been made by a number of botan-
ists and ecologists, such as Transeau (1903), Rigg (1916),
Gates (1926, 1942), Wilson and Potder (19#3), Dutro and
Cohoe (1938), Olson (1943), among others.

Livingston (1905) recognized three types of vegeta-
tion to be distinguished on the lowland in Roscommon and
Crawford Counties, Michigan. They are: (1)0pen-meadow type,
(2)Tamarack-Northern White Cedar swamp, and (3)Mixed swamp.
He also pointed out that the swamps are generally in much
more nearly their original condition than are the upland
forests due to the fact that swamps have not been subjected
to burning nearly so often as the uplands. As to the factors
of distribution of vegetation in the lowlands, he explained:

" ----- but the mixed type (of both coniferous

and hardwood species) is always found on the better
drained portions, where there are humocks raised
out of the saturated soil, and the general level

is a few inches higher. Often this better drainage
seems to come about merely by accumulation of ve-
getable debris, a fact which suggests that perhaps

in time the conifer swamp might give way to the
mixedJ and at last possibly to the hardwood upland

type.

In another paper, "The distribution of the plant

societies of Kent County, Michigan, Livingston (1901) stated

17
that probably the main factor in determining the distribu-
tion of lowland societies is water. However, he stated

further:

' ----- the amount of water is practically the

same in an undrained and in a drained swamp and on
a brook margin; yet the floras are dissimilar,
especially the first two. It has been suggested
that the great amount of organic materials in the
solution of the undrained swamp may effect the
plants physically or chemically and thus exclude
those which occur in the drained swamp. ---- There
remains the other suggestion that the undrained
swamp owes the peculiar character of its flora to
the chemical nature of the soil solution. It may
be lack of oxygen in the soil which shuts out the
plants of the drained swamp."

‘\

Davis (1906) has ably discussed the peat and the
occurring vegetation. He has defined and distinquished the
differences between the "bog", the "marsh“, and the "swamp";
and also grouped the peat lands into eight groups: (l)Elm -
and black ash swamps, (2)Tamarack swamps, marshes, and bogs,
(3)0edar swamps, (5)5pruce swamps, (5)Willow and alder swamps
or marsh, (6)Heath swamps, marshes, or bogs, (7)Grass and
sedge marshes and bogs, and (8)Rush marshes.

Harper (1918) made a study on the plant population
of northern lower Michigan. He believed that some of the
imagined successions on the poorly-drained areas can never
take place without profound topographic changes, which may
or may not come to pass. In illustrating his point, he made

the following statement:

18

"Two genuine types of succession can be studied
to advantage in this region. The first is that
connected with the filling of lakes, etc. with
vegetation and the gradual accumulation of peat
and humus. ----- In such humus grow many plants
which are equally characteristic of the upland
hardwood forests, and this has led some to believe
that the swamps, barring human interference and
unforeseen complications, will ultimately be re-
placed by beech-maple-hemlock forests. But there
are quite a number of plants in this region which
seem to demand both humus and access to mineral
soil or alkaline peat such as Acer saccharum,Tsuga,
Fatus, Eilig, glmps, Quercus alb , Viburnum seep;-
felium etc., and we have no evidence that these
will ever grow on top of deep sour peat. Furthermore,
the swamps are colder at night than the adjoining
slepes, and lack some of the characteristic soil
fauna of clayey uplands, and these foundamental
differences can hardly be obliterated by succession."

 

Gates (1926) has made a study on the plant succes-
sions about Douglas Lake in Cheboygan County, Michigan. He
stated that cedar bog forest will be the terminal stage be-
fore reaching the upland type, whereas the lowland forest of
ash, elm, and red maple is the stage next to dryland on land
which is subject to overflow orz'ISpring flooding.

In 1942, Gates made another investigation on the bogs
of northern lower Michigan. He defined the bog as "an area
vegetated by a flora in which peat-forming types of plants
(including certain herbaceous, ericacious shrubs, and coni-
ferous trees) are particularly abundant." In his paper,
trends of succession are diagramed and the characteristic
vegetation of each stages also are listed.

Clements (1936) has stated that the bogs and muskegs

I“
In ..s:

sorQsAJ-tlca

a: a:
at a.
H. We
c AL.
't not

19
are the prisere subclimaxcs of the boreal and lake forests.

In discussing subclimax, he preposed the term "serclimax"

‘

and made the following statement:

"Serclimax is suggested with a meaning of a seral
community usually one or two stages before the sub-
climax, which persists for such a period as to re—
semble the climax in this one respect.

For the most part, serclimaxes are found in
standing water or in saturated soils as a consequence
of imperfect drainage. The universal sample is the
reed-swamp with one or more of several consocies,
such as Scir us, T ha, Zizania, Phragmites, and
Glyceria. ----- In boreal and subalpine districts
the distinctive serclimax is the peat bog, moor or
muskeg, more or less regularly associated with the
other seral communities of Carex and usually of Larix
and Piceg also in the prOper region."

Potzger (1934, 1941, 1943, 1946, 1947, and 1948) has
made a number of studies on the history and succession of
vegetation in Michigan, principally on the bogs and lowland
forests, by means of pollen analysis. He considered the bog
forests of Apigg, figggg_and ngig_ato be: relics of the boreal
forest caused by the retreat of glaciers.

In "The history of the vegetation of Michigan",
Potzger (1947) pointed out the existance of a "tension zone"
in central lower Michigan, between oak forests of the south
and the coniferous forests of the north.

Nichols (1935), in discussing the physiOgraphic
climax forests in the “Eastern Hemlock" region, has recog-
nized the poorly-drained swamp coniferous forests of balsam

fir, black spruce, white spruce (Picea glauca), northern white

fr!

20
cedar, and tamarack, in varying admixtures, as the physio-
graphic climax types.

Bowman (1944) pointed out that 26 percent of the
forest land in Michigan has been classified by the Forest
Survey as either coniferous swamps or spruce-balsam fir
land. He stated that black Spruce is usually found on the
swamp or bog sites frequently associates with tamarack where
as it may be accompanied by northern white cedar on better
sites. As for the northern white cedar, he claimed that it
is unable to withstand great soil acidity but can endure
shade and a surplus of soil water quite as well as black
spruce; and when conditions are favorable, it will surpass
black spruce and all other swamp species in reproducing it-
self.

Swamp and bog floras have been listed and discussed
by Darlington (1945). He pointed out that there are numerous
swamps in northern lower Michigan, often covered with dense
stands of northern white cedar, tamarack, and black spruce
as well as some bogs with shrub growth.

Braun (1950) has discussed the succession of various
forest communities to one another in sequences of decreasing
water requirements from the bog to the highest ridges of the
forests in northern lower Michigan. She stated that the re-
placement of bog or transitional communities by hardwood
forest can not take place until the swamps are filled or

drained.

In dis
stat of the ‘3
annuities w
out the gener
10.31% 1:03 --
We“; and s
are Lari: a:

N“.

Iii t TIT-l la

N.
in lower l-Iich
the M was
When the 0th
the FNSERCQ
i
Sign: trees 1:

She a

21
In discussing the bog communities Braun said that
most of the bogs and swamps are occupied by coniferous
communities with a few are hardwood swamps. She pointed
out the general sequence of vegetational stages of a deve-
10ping bog -—-- aquatic, sedge mat, bog shrub, and beg
forest; and stated that the principal trees of bog forests
are Larix laricina, Picea maggana, and Thuja occidentalis,
and the ThuJa bog is the ultimate in bog forest development
in lower Michigan. She further mentioned that in many cases
the ThuJa was wanting from the Thugs-lariz-Picea association
where the other two were present in about equal number, with
the presence of balsam fir, white spruce, black ash, white
elm, balsam poplar, red maple and a few other species of
swamp trees to give a mixed character to the type.
She also discussed the river swamp communities in
lower Michigan and stated:
"Shrubby species of willow, alder and spiraea
form a shrub border, white elm, black ash and some
times silver maple dominate in the swamp forest
' strip. The royal fern (Osmunda regalgg) is a fre-
quent subdominant. 0n better drained stream margins,
a few additional hardwood species may enter the elm-
ash communities, suggesting its replacement under
good drainage condition by meBOphytic hardwood forest."
The peat bOgs of eastern North America have been
studied by Dansereau and Segadas—Vianna (1952). In the first
of their papers, "Structure and evolution of vegetation",

‘

succession on the begs has been well illustrated.

Pores

fiscuseed by

 

1929). or t
to the hydra?
begs of 09‘1”
‘ Four

'Lcrthern f0?
been recogniz

if»: others. 1

‘

AI
‘6'

“
~
‘0‘-

;:re or mixed
northern white
Paper birch.

on acid peat i

2 0301.1
tier

White as

‘. A . ‘
.mlace tamara

HM
wk nah, re:
soI‘Etimes has

'9“
' ‘h
“a;

+
«fins itsc

1:81: 1
“13 found

in:
not strom

3.Ta:a

“‘3 a
”Amati. .: .

lam

"em white

22
Forest succession in northern Minnesota has been
discussed by Bergman and Stallard (1916), and by Stallard
(1929). Of these, parts of the papers have been devoted
to the rhydrarch succession. . Likewise, plant succession in
bags of- central‘Minnesota‘has been studied by Conway (1949).
‘ Four distinguished forest types on wet sites in
"northern forest region" of the eastern United States have
been recognized by the Society of American Foresters (Hawley
and others, 1932). They are:
l.Black spruce type ----- composed of black spruce,
Pure or mixed with a minor portion of balsam fir, tamarack,
northern white cedar, black ash, red maple and occasionally
Paper birch. It is a subclimax type to be found in swamps,
on acid peat with little or no drainage.
2.Northern white cedar type ----- composed of nor-
thern white cedar, pure or predominating. Associates may
include tamarack, balsam fir, yellow birch, paper birch,
black ash, red maple, black spruce, white pine, and hemlock.
s°metimeshas an under growth of. alder. If undisturbed, it
maintains itself as long as the swamp remains wet. It is
“anally found on sites with slow drainage, high water-table,
‘nd not strongly acid.
3.Tamarack type ----- composed of tamarack pure or
I:‘I‘Odominating. Common associates either black spruce or

11c’I'thern white cedar or less commonly both.0ther associates,

23

mostly, subordinate, include red maple, black ash, and

aspen. It) is chiefly found in peat swamps with little or no
drainage, and is succeeded by black spruce on undrained acid

post. or by northern white cedar in better drained less acid

4.Black ash-American elm-red maple type ----- Black

33h occurs least often in other types and therefore may be

cons idered as an indicator species for this type. It is a

Climax for the site. In Lake States, the associates include

balsam peplar, balsam fir, yellow birch, and less commonly
‘fllflste pine, tamarack, northern white cedar, basswood, etc..
It occupies moist to wet muck or shallow peat soils in swamps,
gull-lies, and small depressions of slow drainage or along
alugegish streams. .

Gates (1930) has made a study on the aspen associa-
tion in northern lower Michigan. He pointed out the occur-
zwhee of aspen association is the result of virgin or second
8I'GWNth forest from burning.- In his paper, he recognized
three types within the aspen association : (1)10w1and type,
(2)3andy upland type,and (3)01ayey upland type. For the

1°Vland type, he stated:

"The sandy lowland types of aspen association
is distinguished by the dominance of Pepulgs pgge
m loide , and it will be replaced by either Thuja
association, or a mixture of spruce-balsam, or the
lowland forest of black ash and American elm."

‘

C)

 

 

ESCRIPTION OF THE REGION

A.History and general description

Ecologically speaking, the Higgins Lake area is
located within the Hemlock-White fine-Northern Hardwoods
Formation which lies north of the tension zone between this
Formation and the Deciduous Formation of the south in the
State of Michigan (Fig. 1).

Geographically, the area is located in the northern
Central part of the lower peninsula of urchigmn,and is with-
in the scepe of the Higgins Lake State Forest which is com-
pOsed of parts of four Counties: Missaukee, Kalkaska, Craw-
1‘0rd, and Roscommon (Fig. 2). The Higgins Lake State Forest

encloses an area of 311,480 acres within its boundary, and

has a not state ownership of 195 ,475 acres (Michigan Depart
ment of Conservation Biennial Report, 1951-1952).

In 1899, a State Forestry Commission was authorized
to study the forest situation in Michigan, and set up re-
aGloves but nothing was accomplished until 1903. That year,
two "Reserves“, the Higgins Lake Reserve in the northern
half of T2411, R41! and the southern half of T25N, R4W, Craw-
f.‘Drd County, and the Houghton Lake Reserve in T21N, R3W and
T3111, Rim, Roscommon County tOgether comprising 35,000 acres

°1' tax-reverted, cutover and burned over land were set aside.

 

 

 

 

 

 

    
 

BROWN
FOREST SOIL

.....___. Oak-hickory White pine

 

...—...,Northern boundary of Beech-Maple region

Fig. 1. Map of Lower Michigan showing the location of
the Higgins Lake area in relationship to the
Hemlock—White Pine-Northern Hardwoods region
and also its relation to some of the tree
ranges and soils. (After Braun, 1950, and
Veatch, 1932)

26

'\

Oflo_.:_.:_oT .floflowo—Oq

.Lo‘"

0
O O
o—g-—-..-—-o—_.

 

¢_o.——.—O—Q—.
.
o

 

 

 

. :Mnsspsuxsa. . ! RDSCONMQN
'. I I i

L__..L..__L.__.;_H._iu__.;_.__J.__uL_..

.__.:_._:_.._...:__ 'T

' o
. O
0
.fl.;. —..——.—'

LEGEND

—- - -—- coumv BIN-Mosaics

TOWNSHIP MNDARIES

souuomv or-Hmnws LAKfi STATE rosesr

LAKES

 

Fig. 2. Map of Higgins Lake State Forest showing
its boundary within the four adjoining
Counties: Missaukee, Kalkaska, Crawford,
and Roscommon. (After the State Owner-
ship Maps of the four Counties, Michigan
Department of Conservation, June, 1951.)

Ecuever, no act!
like Reserve un‘.
Forest was th
active manageme:
The out.
it the time of i
5H dense fore:
“'3 3039 Open 1;
11515th of pin

L.“ .
at. iw 30.: 3 3‘50“: ,-

27

However, no activities were being carried out on the Houghton
Lake Reserve until 1911, therefore the Higgins Lake State
Forest was the first of the State Forests to be put under
active management.

The entire land area of the Higgins Lake State Forest,
at the time of its first occupation by white men, was covered
by a dense forest except for a small acreage of bog or marsh
and some open land in the drier sandy plains. The lumber
industry of pine on large scale started about 1870 and of
hardwoods about 1900. At the present time, the original forest
has long been cut ovor except in a very few small areas and
"00d1ots, and may be a few places in swamps.

Several types of forest were represented in the virgin
f0Pest as recorded by the Soil Economic Survey: (l)The hard-
w‘Mbd forests , in which sugar maple and beech were the dominant
aDefines, and elm, ash, basswood , yellow birch, and hemlock
Wer‘e the subdcminantespecies; (2)the mixed conifer and hard-
w°cd forests, in which such species as elm, ash, red maple,
8When, and yellow birch were intimately associated with white
pine, hemlock, spruce, and fir; (3)the pine forests, in which
“hikte pine, red pine and Norway pine or red and Jack pine
DI‘edcminated; and (4)the swamp forests, in which the dominant
ape<‘:ies were northern white cedar, spruce, balsam fir, and

taslimrack. Much of the forest land is at present covered with

a flame

Der
of pet

the nc

II”,
L. the

‘
a:
No 1.1
. h. .
H .4 o‘
.Vlo 5 .

itle

V

‘ .

o»
n. .

... x r! . I

a . in» t .
‘ C a“ o‘ I \
3‘ tr ... F.“ PH

0% O .3 at t D. t
..
C
.
'
O\
.‘

28

a dense growth of brush, briers, and grass or grown up to
ocrubby aspen, oak, and rod maple, with little natural
reproduction of the original dominant species.

In the early days of the area's development the
lumber industry unquestionably brought the largest number
of people into the area. During this period of the late
seventies the railroads were extending their lines into
the northern regions of the state and opening the way for
further settlement in this area. At the same time agricul-
ture was also gaining a foothold so that the population on
farms was being increased. Between 1900 and 1910 the rate
of farm settlement was considerably increased by tho acti-
vities of outside real estate companies that sold cut-over
lands to now settlers.

with the decline in the lumber industry after the
excessive cutting, the farmers lost an important and easily
accessible market for their produce and surplus labor. With-
out this market, many farmers were forced to leave their
farms. Extensive abandonment has since occurred on the lands
of low productivity and least favorable locations which were
chiefly those lands sold by the previously stated companies.
The tracts and lots which have been abandoned by their owners
now make up a large part of the present area of the Higgins

Lake State Forest.

The recs
resort and recre
roads Ind later
Emu to attract
tremenieus numb-3
:ghtOI Heights
others. Higgins

rams hunting ,

29

The recent development in this area has centered on
resort and recreational propertya‘With the advent of improved
roads and motor transportation, Houghton andHiggins lakes‘
began to attract more visitors each year. There are now a
tremendous numbers of dwellings in the resort villages of
Houghton Heights, HoughtonLake Village, Prudenville and
others. Higgins and Houghton Lakes have since become a

famous hunting, fishing and outing ground.

30
B.Clinate

Michigan is located in the heart of the Great Lakes
region and has the longest shore line in the States. The
large bodies of water tend to equalize the nearby land tem-
peratures and this is especially true of lower Michigan,
where the effect of cold waves sweeping down from the north-
west is modified by the warmer water of the Great Lakes.
This lakes for lower Michigan a more equal and less extreme
climate than occurs in the states of similar latitude on the
other side of Lake Michigan.

In the Higgins Lake area, the climate alternates
between continental and semi-marine conditions. The marine
type is due to the influence of the lakes and governed by
the force and direction of the wind. The weather becomes
continental in character when there is little or no wind,
and a strong wind from the lakes will transfer it into a
semi-marine type.

The winters are long and rigorous in this area, as
temperature below freezing prevail from November to March,
inclusive; and occasional freezes may well be expected in
September and May, or even in June. This limits the frost
free or the growing season to a comparatove short period.
The average period for the region as a whole is about 105

days. It is derived from the data of six weather stations

in and near the
SE to 125 days .
W195 days at R
presence of larz
ms.

The main
W1 telperatu
he 15°F” and
precipitation (1
"m8! snowfall
1" Percentage s
“d and “More

Tl“ Pr‘ec
through.“ the 3

six-loath De 1‘ ice?

31
in and near the Higgins Lake State Forest which range from
88 to 125 days. The low of 88 days at Houghton Lake Station
and 95 days at Roscommon Station are probably due to the
presence of large bodies of lakes and swamp lands in those
areas. .

The main climatic features of this region are a mean
annual temperature of about 43°F. (average January tempera-
ture 18°F., and average July temperature of 68'F.); a normal
precipitation (including melted snow) of about 29 inches; an
average snowfall of approximately 70 inches} low evaporation;
low percentage of possible sunshine; and rigorous winters
and mild summers.

The precipitation is distributed relatively even
throughout the year with a slightly greater amount for the
six-month period from May to October. Most of the precipita-
tion occurs as slow rains with rare occasionsof destructive
downpours. Snow ordinarily forms a permanent ground cover
from November to March and serves to prevent the soil from
freezing to a great depth. The prevailing winds are westerly
and rarely attain high velocity to cause any destructive
damages.

Weather data of this region are presented in Table l.
The data are adapted from "Climate of Michigan" in Climate
And Man (U.S.D.A. yearbook, 1941), and expanded by using

Climatological Data for Michigan, U. S. Department of Commerce,

Heather 3.:

Stations w

 

32
Weather Bureau, from 1936 to 1952. The locations of Weather

Stations which are near.the Higgins Lake are shown in Fig. 3.

Table 1. Weather data of the Higgins Lake area..

 

 

 

 

 

Temperature _F. Killing Frost Ave.Dates'Precipi-
Jan. July ~ Last in First in tatiog
Station 1 ave. ave. 4a b 9:1, Spring; Fall 2 1 Inches
Higgins .
Lake 50 17.7 66.9 106 —39 17 June 6 Sept.l6 102 48 29.39
Houghton
Lake 36 19.8 67.4 107 —48 36 June 10 Sept. 6 88 37 27.40
Roscommon“ 8 18.6 67.3 ------ 8 June 2 Sept. 5 ‘95 7 29.98

Graylinsfl 51 17.7 67.6 106 -41 54 May 29 Sept.18 112 53 31.07

Kalkaska '22 17.6 67.8 106 -34 24 May 23 Sept.25 125 35 30.20

Lake City 30 19.3 68.6 106 -41 30 May 728 Sept.2l 116 29 26.44
*Data compiled from Climatological Data for Michigan,

U.S. Department of Commerce, Weather Bureau, from
1939 to 1946 only.

 

 

 

 

 

 

 

l ----- Length of record years
a ----- Maximum temperature recorded
b ----- Minimum temperature recorded
2 ----- Length of growing season in days
Precipitation in Inches
Stat o J . Feb. Mar. App. May June July Aug. Sept Oct. Nov. Dec.
Higgins
Lake 36 1.46 1.47 1.82 2.44 2.79 2.59 2.68 2.90 3.21 2.65 2.29 1.62
Houghton

Lake 24 1.34 1.21 1.99 2.53 2.74 2.77 2.38 2.73 2.92 2.97 2.37 1.45
Graylins 39 1.56 1.52 1.70 2.36 3.37 2.89 3.41 2.90 3.43 3.16 2.73 1.70

E:1kaska 23 1.49 1.23 1.94 2.20 2.94 2.60 3.07 3.19 3.30 2.87 2.40 1.95
ke

0111. 11 1.47 1965 1.50 1.96 2.52 2.65 2.30 2.25 2.94 2.33 1.79 1.29
l ----- Length of record in years

n
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P
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g 1 o
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.5
.12
LIL-1 7
O
.3
o6
* f ”L
1 Grayling 2 Higgins Lake 3 Houghton Lake
4 Roscommon~ 5 Kalkaska 6 Lake City

7Rose City (Lupton)

Fig. 3. Map of Lower Michigan showing the location of
U. S. Weather Bureau Stations in and near the

Higgins Lake area whose weather data are used
in this study.

33

.._«

”I

as.

-~

34

C.Geology

The surface features of the southern peninsula of
Michigan are product of glaciation. The glacial drift, which
covers so deeply much of the rock surface of the southern
peninsula,was brought in largely by an ice sheet or conti-
nental glacier which.moved southeastward from the highlands
of Canada. According to Leverett (1912), " ----- the average
thickness of the drift in the southern peninsula‘is.ab0ut —
300 feet, wheeeas in the high interior of the north part of
the peninsula may.have over 1000 feet."

In the Higgins Lake area, the surface features have
been changed very little since the close of the last period
of glaciation. Much of the land formations in this region
is level or gently rolling which can be grouped into three
physiographic divisions; (l)The outwash plains, (2)the till
plains, and (3)the moraines. The underlying bedrock has been
covered to a considerable depth by glacial drift and outwash
so that no surface outcroppings of hardrock occur. According
to Michigan Geological Survey, it is 417 feet at Roscommon
in Roscemmon County. The structure of the drift is rather
variable in Michigan. Leverett (1904) stated that:

" ----- Sand and gravel form a notable part of

the drift material and much of the till is loose
textured. This great amount of loose textured
drift seems attributable to the excessive glacial
drainage resulting from the convergence of the
ice lobes."

35

Surface land formations of the Higgins Lake State
Forest are shown in Fig. 4. The outwash plains are mainly
level while the smooth rolling uplands occupying the till
plains and the moraines make up the rolling to hilly up-
lands. Considerable areas of swamp and wet lands occur
around the lakes and along the rivers, chiefly on the out-
wash plains; and also some of the swamp soils which are
scattered in spots occur in the deeply cut valleys and on
the rolling to hilly uplands.

A table-land ranging in height from 1,200 to 1,400
feet which 'embraces‘w“ parts of Crawford, Kalkaska, Mis-
saukee, and Roscommon Counties to'form a divide between three
large river systems. As a result, three drainage basins
are formed as shown in Fig. 5. In the Higgins Lake State
Forest region, areagin Crawford County and roughly the
eastern one third of the area in Roscommon County drain
into the AuSable River, which flows eastward to Lake Huron.
The area in Kalkaska County and approximately the northern
one third of the area in Missaukeo County drain into the
Manisteo River; while the rest pf the area drains into the
Muskegon River which heads in Higgins and Houghton Lakes.
Both the Manistee and Muskegon Rivers flow westward‘toilake,
Michigan.

The abundance of lakes in the southern penisula is

an indication of the imcomploteness of the drainage, There

‘ .
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36

 

 

 

 

 

1.3] 4.~‘\‘\‘\\‘\ \\

w$§§§

 

      
   

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< moxx
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.0593: \\\\\\§ \ sou—1H U
25.8 2:. W 3.25 \YX!

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. . 33m 83 can;

 

g

 

 

mxsm

  
 

 

37

 

Might

 

/.__-__-__..____,__-

m MUSKEGON AUSABLE

 

 

Fig. 5 Map of Lower Michigan showing drainage

basins

serving the Higgins Lake area.

38

are two large lakes and a few smaller ones in the Higgins
Lake area. Houghton Lake.is the largest inland lake in the
State of Michigan. It has an area of approximate thirty-one
square miles and is more than eight miles long and over four
miles wide. It is a shallow lake which does not exceed 25
feet in depth, and has heavy growth of vegetation. In con-
trast, Higgins Lake, which has an area of about 15 square :
miles is deep and is fed by springs. It also isslocated on a
higher elevation, 1,160 feet above the sea level, as compared
to 1,125 feet of Houghton Lake.

The general elevation of the region ranges from 1,000
to 1,400 feet above the sea level or about 400 to 800 feet
above the level of Lake Michigan and Lake Huron. Elevations
of the level plains vary from 1,000 to 1,200 feet, while the
hilly lands usually reach an elevation of 1,150 to 1,300 feet

and may reach as much as 1,400 feet at places.

 

 

v
.4
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o
a"
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1
a.

D.Soil

The soils of Michigan are developed from the materials
deposited by repeated glaciers which covered the state at the
close of the Tertiary. This glacial drift, which covers so
deeply much of the rock surface of the lower peninsula has an
average thickness of about 300 to 600 feet (Leverett, 1915).

The soil conditions are chiefly determined by the sur-
face geology and the topography. The original character of
the soil, whether rock, sand, clay or marl, depends upon the
geological relations. From the vegetation standpoint the to-
pographic relations are much more important, since they con-
dition the presence or absence of drainage and cause variations
in aeration and humus contents.

The soils in the Higgins Lake region can be classified
into two major divisions: Those deve10ped under conditions of
good drainage and medium moisture and those developed under
poor drainage or excessive moisture. In this paper only that
of the latter are to be discussed.

Poorly-drained areas or the swamp land makes up a
considerably large portion of the land area in the northern
counties of lower Michigan. The soils develOped under condi-
tions of poor drainage in the area of this study are grouped
into three series. They are the Newton soils, the Bergland
soils, and the organic soils. General descriptions of these

soils are as follows:

40

l.Newton soils

These soils occur on poorly-drained sand plains, on
the flat wet borders of shallow lakes and swamps.They consist
of a dark layer of loamy sand or sandy loam soil underlain by
gray or dingy-white wet sand. The dark color, due to organic
matter accumulated under wet conditions extends to a depth
ranging from 3 to 15 inches. Most of these soils are medium
or strongly acid in reaction.

The original tree growth consists mainly of white
pine (2iggg,ltrobus) with more or less northern white cedar
(Iggjg gccidentalis), black spruce (Picea mariana), and balsam

 

fir (Agigg‘balsamea) (Soil Survey of Roscommon County, Michi-
gan, 1924). At the present time, most of the land is covered
with a dense growth of alder, willow, and aspen.
2.Bergland soils

These are the heavier mineral soils which have devo-
leped under conditions of poor drainage. They occur only in
small areas as swamp lands in swales and on the borders: of
peat and muck swamps. These soils are nearly neutral in re-
action. The surface soil is dark gray or nearly black loam
or clay loam with high content of organic matter. It is under
lain by a gray or drab plastic clayey layer, mottled with
yellowish or rust-colored spots showing the typical charac-
teristics common to clay soils existing under permanent wet

conditions.

1

¢

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41

The native vegetation consists mainly of elm and ash,
with smaller amount of red maple (Acer rubggg), basswood
(Tilig amegicana), spruce, balsam fir, northern white cedar,
hemlock (Egggg canadensis), and white pine.

3.0rganic soils

The organic soils are composed dominantly of plant
remains, and constitute a distinct class in comparison with
soils which are composed principally of mineral or inorganic
matter. In this region, the organic soils occur in swamps,
bogs, and marshes. The deposits have accumulated in permanent
wet situations such as in irregular shaped flat areas where
the under drainage is obstructed, in stream valleys, and in
certain types of lakes, some of which have been completely
filled by vegetation. The organic deposits range from one
foot to as much as 40 feet. Based upon the vegetational ori-
gin of the organic deposits, and the degree of its decomposi-
tion, the organicsoils in this region are further grouped
into the following types:

a.Riflo peat
Rifle peat is brown or dark brown, coarse, woody or
loamy muck or peat very rich in organic matter, which is
underlain at a depth ranging from six to 20 inches by fibrous
or coarser textured plant material with very little decomposi-
tion. The average depth of the water-table is about 10 to
20 inches. This organic soil is acid or nearly neutral in

reaction. It occupies the largest aggregate acreage of the

 

 

 

amp land in t:
The veg;
northern white
its; Elsi;
Whom the swamp

Brown up in dens

birch, whereas t

 

Erowth of sedges
b.6reenwood
This is 1‘

mt in which th
even at the surf
“id in reaction

Except, during ve‘

H'uCh of

 

Subs tut“ . T L.

 

 

42

swamp land in this region.

The vegetation consists of a dense swamp growth of
northern white cedar, black spruce, balsam fir, and tamarack
(L§§;;,lar1cina), with occasional hemlock and white pine.
Where the swamp land has been logged and burned over it has
grown up in dense thickets of aspen, alder, willow, and
birch, whereas treeless open areas are covered with a heavy
growth of sedges and bluejoint (gglamagrostig canadenglg).

b.Greenwood peat
This is fibrous or coarse-textured, loose, brownish
peat in which the plant material is only slightly decomposed,
even at the surface. This kind of peat is uniformly strongly
acid in reaction. The water—table is at or near the surface,
except during very dry periods.

Much of the Greenwood peat land has a semi-floating
substratum. The Open heath bogs support leatherleaf (Chamae-
daphne calygglata), labrador tea (LQQEQ groeglandicum), blue-
berry (Vggcinium sp.) and Sphagnum moss (Sphagnum sp.) with
occasional growth of tamarack and black spruce (Soil Survey
of Roscommon County, Michigan, 1924). Some of the more open

land is marsh grown up chiefly to sedges.

43

E.Covor type

In the northern counties of Michigan, swamp land
makes up a considerable portion of the land area. Most of
the timberod swamp lands at the present time are occupied
by coniferous swamp species; the rest of the areas are occu—
pied by the swamp hardwood species and in some open places,
marshes and bogs.

The swamp forest growth varies considerably in age
and size. Many areas have been closely cut over and are now
occupied by dense reproduction. Other areas are well along
the second growth stage and a few lightly out or virgin
stands still remain. Generally speaking, the swamp stands
are nearly all well stocked. This may trace_back to the fact
that.the swamps are usually wet, their cover has suffered
less from fire than that of the uplands.

The poorly-drained or swamp land in the Higgins Lake
area can be grouped into the following vegetation types
(Land Economic Survey Reports of Roscommon, Crawford, and
Kalkaska Counties, Michigan.):

l.Swamp Hardwood type

The major species of this type are elms, black ash
(W gm), red maple, and yellow birch (m M).
On the better-drained areas, aspen, white birch (Betulgppgpze
rifogg), and alder may enter, while poorer drained sites

include so
mi tamara
lowlands t
Q‘dently wh.
8011. The
“fining 1
areas of 3‘
types of PE

44

include some balsam fir, black spruce, northern white cedar,
and tamarack.

The Swamp Hardwood type is usually found on areas of
lowlands that possess a very moist mineral soil or more fre-
quently where a shallow muck layer overlies a wet sandy sub-
soil. The usual location for such site is the swamp margins
bordering the highlands. It also occurs commonly on those
areas of swamp land that are occupied by better decomposed
types of peat and muck of relatively shallow depth on a mine-
ral substratum. ‘

The Swamp Hardwood type usually exists as an uneven
aged stand and rarely attains the volume produced by the up-
land hardwood type.

2.8wamp Conifer type

Northern white cedar, black spruce, balsam fir, and
tamarack predominate the Swamp Conifer type with the associa-
ted species of red maple, yellow birch, elm, black ash, and
aspen. g

This mixture of species will vary in composition with
the site on which the stand is found. 0n better drained sites
balsam fir, northern white cedar, red maple, and yellow birch
predominate, while black spruce and tamarack become more pro-
minant on the poorly drained sites. Between these two extremes
are found many variations in composition depending on drainage,

depth of the peat, acidity, and soil temperature.

45

The majority of the stands have been cut over, or at
least culled over once or several times. Cut-over areas re-
produce themselves woll, while the severely burned over areas
are occupied by a dense growth of alder and willow which
offer much opposition to the coniferous reproduction.

3.Lew1and Aspen type

Aspen stand is found to occur on nearly every class
of soils and site, from the denuded hardwood and pine uplands
down to the swamp lands.‘ On most of the soils it exists as
a temporary cover and is quite easily replaced by other species
1h gimo,fllHowever,*repeatedlyv' burned over areas generally
restock the aspen type because of its ability to reproduce
from sprouts. .

Trembling aspen (Pogglus tremuloides) is the major
species in the Lowland Aspen type. The most important inva-
ders are northern white cedar, balsanlfir and spruce, black
ash and American elm (Egggg americana).

4.3wamp Shrubs of §2;;ngornus-§;ggg type

This type consists of a mixture of §g;;g, Cornus, and
A;ggg.orranyIOne species alone. It occurs frequently on swampy
or poorly drained areas which have been subjected to repeated
burning, in addition to be a common associate in the Swamp
Hardwood and Swamp Conifer types. It is also the principal

growth along narrow stream bottoms.

...a

poorly-‘

mars

of

"
1‘.

ten

111.”? OH '

I
a

I‘

H

the n

llow

I
a

i

c

46

5.Marsh and Bog types

These are the vegetation types which occur on open
poorly-drained land.

a.Marsh

In this region, there is one considerably large area
of marsh, among other places, located just west of the Hough-
ton Lako along Highway US-27.

These marshes occupy the low flat lands which are
wet but have a surface drainage outlet, principally along
narrow winding streams.

Sedges (Egggg), bluejoint. and other grasses form
the principal vegetation. The wettest sites usually contain
some cattails (Typllg 2511391194) while the relatively better
drained areas have more lgraSs;.i . A patchy growth of
.willow (§§;;§ sp.) is often associated with this type.

b.Bog

This type occurs on the bog areas and has a growth
that consists mainly of leatherleaf, with a mixture of
Sphagnum moss, labradorrtea, and Iaccinium.

These bogs are usually treeless but in case the
drainage is somewhat better, an Open stand of black spruce

or tamarack may be present.

VEGETATION ANALXSIS

A.Field procedure
l.The field work was conducted during the summer of 1952.
A preliminary reconaissance was made to investigate the areal
extent, gross aspects of vegetation cover, associated soils,
tepography, and other local conditions of the entire region.
Typically representative stands were chosen as sample
plots based on the vegetation and soil type and the considera-
tion that they had the least disturbances either due to na—
tural or human causes, such as fire, destructive cutting, etc.
_ Five types of vegetation were recognized on the poorly
drained soils in the Higgins Lake area:
a.The Swamp Hardwoods of Elm—Black Ash-Red Maple Associ-
ation.
b.The Swamp Conifers sf Black Spruce-Balsam Fir-Northern.
White Cedar Association.
c.The Lowland Aspen of Populus-§§;;;_Association
d.The Swamp Shrubs of Salig-Cornus-élgug.Association.
c.The Marsh or Open-Meadow Association.
2.3ampling method
Quadrats_were used in sampling the vegetation in
order to obtain maximum accuracy. The quadrats were placed

in each vegetation type according to the various successional

48

stages. This would probably give a better interpretation for
the objective of this study, forest succession.

Sample plots were established depending upon the ex-
tent and configuration of the stand, and are laid in such a
way so as to avoid the border effect. They were being dis-
persed and randomly distributed in the entire region. Iow-
ever, stands which were located too deep‘in the swamps, too re-
mote from the roads, or too widely scattered were avoided so
as to reduce much of the travelling time especially during
the period of collecting instrumental data.

The number of sample plots in each vegetation type
were determined by the homogenity of stand composition, the
extensiveness of the type refering to the region as a whole,

and in some instances, the accessibility of the stands-':

Nested quadrats were used in this study. The sizes
of different quadrats were: -
a. 1/5 acre quadrat for tree species more than six feet
tall, and shrub species more than 25 feet tall.
b. l/lO acre quadrat for shrub species or any vegetation
from 6 to 25 feet tall.
c. l/lOOO or milacre quadrat for all vegetation below
six feet tall.
This classification is a common practice in forestry

research work. The principle is to select the right size plot

 

49
which is small ornugh to work with, and fairly effective
in representing the pepulation and to achieve the objective
of the study.
Individual r. tree species, on 1/5 acre plots, were
tallied and recorded according to the size classes desig-
nated by Weaver and Clements (1938), with a slight adjust-
ment in the size class 2:
Size class 2. Medium reproduction ----- tree species
more than 6 feet tall or up to 0.9 inches 0.3.3..
Size class 3. Large reproduction ----- 1 inch to 3.5
inches D.B.H..
Size class 4. Small trees ----- 3.6 to 9.5 inches D.B.H..
Size class 5. Large trees ----- 9.6 inches or more D.B.H..
Individual shrub species on 1/lO acre plots were
tallied according to Height classes arbitrarily set up by
the author based on the natural height growth of the shrubs
in the region. Two classes Iere used in this study:
Height class 1. Vegetation from 6 to 15 feet tall.
Height class 2. .Vegetation over 15 feet and up to 25 ft..
In the milacre plots, all vegetation was recorded by 1
classes arbitrarily designated by height in order to show
their stratification as occurring in nature. Three classes
were used:
Class 1. Plants up to 0.9 feet tall.
Class 2. Plants from 1 foot to 3 feet tall.
Class 3. Plants over 3 feet and up to 6 feet tall.

5.9

50
Tree and shrub species were recorded by number of
stems into their corresponding height class in the milacre
plots, with a few exceptions where some of the shrubs were
difficult to count and therefore were recorded by"coverage§
Cayerage in this paper is defined as the ocular estimation‘

of the plants which cover the ground surface in vertical

 

projection as expressed in percentage. All herbaceous plants

W**
1"" t "—

and pteridophytes were tallied by coverage..

51
B.0ffice procedure

..1.Summerization of field data

.a.Concerning tree species as ----- The objectives are to
provide data which upon processing will yield the following
lndices for the tree components of the stand.

1(1)Frequency--- a measure of dispersion eXpressed as If

 

the percentage of quadrats in which: t
.(a)a given species occurs (i.e., frequency per species)

Fq of sp. A = No. of uadrats n which s . A occurs 100
Total no. of quadrats examined

.(b)a given size class of a given species occurs (i.e.,
frequency per size class per species)

F of sp. A = No.ggugd. in which size class n of A occur x 100
Total no. of quadrats examined

i(0)R61&t1ve.froquency--~-~ a relative expression as a
percentage of the frequency value for a given species

and based on the total of the frequency values for

Spec 1‘. s

Fr of sp. A = Freguency of species x 100
Sum of frequency values for all sp.

.(2)Density—--- a quantitative measures of the number of

individuals of a species per unit area.

D of .113. a : Igtaino. of.indiv.iduals‘ orsp. A counted xioo
Total no. of quadrats examined

-(3)Relativegdensity~--- a relative index of plentifulness

expressed as the percentage representation of '

 

52
(a) each species, based on the total number of indivi-
duals counted (i.e., relative density per species)

Drs of sp. A = Totgl no. of individuals ofggp. A counted x100
Total no. of individuals of all species

(t0 each size class of a species, based on the total

number of individuals in that size class (i.e.,

“.-fi

relative density per size class per species) L

Dr of sp. A = Totgl no. of size class n stems of s .A x 00
Total no. of size class n stems of all species

(4)Basa1 area ----- a quantitative measure of dominance as
expressed by summation of the area occupied at breast height
by holes of a given species.

(5)Relative basal area ----- a relative index of dominance
eXpressed as the percentage of basal area occupied by a given
species.

Br of sp. A = Total B.A. occgpied by individualgfiof sp.A x100
' Total B.A. occupied by all species

(5)1mportance value ----- a summation index indicating
overall importance of a given species and calculated by
adding the separate values for relative frequency, relative
density, and relative basal area for that species.

‘Inportance Value of sp. A 8 Fr + Drs + Br
ILConcerning shrub species ----- the following indices are
produced from field data:

LIJFrequencyj

(8) a given species occurs (Fq)

(b) a given height class of a given species occurs (F)
(9) relative frequency (Fr)
(2)Density (D)
(3)Relative density
(a) of each species (Drs)
(b) of each height class of a species (Dr)

(4)Abundance (A) ----- the average number of individuals

'9 ~ ’I

of a species per quadrat considering only quadrats in which

the species is presented.

A of sp. B 8 Total number of individuals of sp. B counted

No. of quadrats in which sp. B occurs

 

But D 3 Total individuals
Total quadrats

And Fq = Quadrats of occurrence_g5100
Total quadrats

Therefore A = lOOD/Fq
(SNRelative abundance (Ar) ----- a relative index of
abundance expressed in percentage

Ar of sp. B 3 Abundance_of sp. B
Abundance of all sp.

¢.Concerning herbaceous species ————— All plants below
6 feet tall were grouped into form groups of Tree, Shrub and
Vine, Fern and Fern-like plants, and Herb. They are listed
by species alphabetically by percentages of coverage accord-

ing to the three classes designated.

.-

54
2.0911ection, identification and nomenclature of the
vegetation
Specimens of all plants found in the quadrats were

collected and deposited in the Herbarium of Michigan State

College.
Nomenclature for the trees and part of the shrubs {I
‘ i

W. ‘
j-

are following the "Check List of Native and NaturalizedrTrees
of the United States (Including Alaska)" (U .s .D mums. Agr.
Handbook No. 41, 1953). All other planta follow Gray's
Manual of Botany (Fernald, 1950).

A check list of the plants encountered in this study
is supplemented in Appendix, listing 171 species of which
there are 20 trees, 51 shrubs, lO pteridOphytes, and 90

herbaceous species.

55

C.Vegetation inventory

As mentioned previously, five different vegetation
types were found on the poorly-drained soils in this region.
vegetation inventory was carried out in 1952, from June to
October.

An increment borer was used in the field to deter—
mine treo age. The largest tree was selected in order to
determine the sequence of ecesis of various species in each
1/5 acre quadrat.

To each of the 1/5 acre quadrats, four milacre plots
were set up to study the ground vegetation. Plants in the

milacre quadrats are grouped into four presence classes as

following:
Abundant ----- Plants present in all four quadrats.
Frequent ----- Plants present in three quadrats.
Common ——————— Plants present in two quadrats.
Rare --------- Plants present in one quadrat.

Summarized data of tree species by size classes, and
glants of the herbaceous layer by height classes, and, if
there is any, the shrub species by height classes are com-

piled and summarized for each vegetation type.

 

0
d
.p ,
0
.~ \
v
I
o
‘ .
-.n. .}
.-efi-v‘ -. ~
.- . --.»... ox
- Y

t-.

56
1.Swamp Hardwoods of Black Ash-American Elm-Red Maple
Association
The principal hardwood species to be found in this
type were the black and green ash, red and silver maple,
American and slippery elm, basswood, and yellow birch. ;
Northern white cedar and balsam fir were the associated {I
coniferous species. Four stands were selected as sample plots: A
a.B1ack Ash-Northern White Cedar-Balsam Fir stand
Location ----- Roscommon County, Markey Township,
mm, R331, 33 1/4, Section 24.
Soil type -§--- Rifle peat.
This stand was a mixture of hardwood and coniferous
species. Black ash was the predominant species which was
associated with northern white cedar and balsam fir. A few
green ash, red maple, yellow birch, hemlock, American elm,
and black spruce complete the tree speCies found-in the
quadrat. Speckled alder (Alggg rugosa) was the only shrub
species present.
The largest tree was a northern white cedar, 13.5
inches D.B.H., 160 years old.- The remaining species of
balsam fir, red maple, green ash, and yellow binch were all
about the same age, around 60 years.
Black ash and balsam fir were reproducing well,
followed by northern white cedar. Reproduction of red maple

was abundant, but almost all were one year Old seedlings.

 

57
Plants of the herbaceous layer were grouped separate-
ly into tree, shrub and vine, fern and fern-like plants, and
herb groups according to the four presence classes, with the
number of tree seedlings also listed:
Tammi

Abundant --- Acer rubrum (299)
Abies balsameg (l8)

 

Frequent --- Fraxinus ni ra (18)
leus rubra (3)
Common ----- Thuja occidentalis (10)

fistula lute; 10

Rare ------- Quercus rubra (l)

 

§hrubs and Vine;

Abundant --- Rubug pubescens

Common ----- Alnus rugosa ‘ Clematis virginiana
, Ilex verticillata Mitchella repgns
Rare ------1 Lonicera canadensis Ribes lacustro

Rubus ideaus

Ferngfiand Fern-like Plants

Common ----- Dryopteris spinulosa Dryepteris cristata
Onoclea sensibilis

Sedgeg, Grasses and Herbs

 

Abundant --- Galina triflorum Carex intumescens

Frequent -—- Agrostis hyemalis Aster lategizlorug
Cornus canadeggis Galium trifidum
Lgcopus americanus Maianthemum canadensis
Mitella nudg

Common ----- ggalia nudicaulis Gjyceria striata
Lisimachia thyrsiflora Viola blends

Rare ------- Aster pgniceus Rannuculus sp.

chtellaria lateriflora
Solidago ulmifolia Thalictrum dagycarpum

“so;

 

 

“ .-
w.
.. ....
- .o _

out .1

nu

58
b.Ash-Red Maple-Elm stand
Location ----- Roscommon County, Lake Township,
T23N, R4w, NW 1/4, Section 3.
Soil type ----- Bergland loam.
This was a well stocked stand composed of black and
green ash, red maple, American and slippery elm as dominant i
species, which associate with northern white cedar, balsam L
fir, and a few basswood. A few shrubs were the Amelanchier
sp. and Corylus cornuta.
The largest tree was an American elm, 20.1 inches
D.B.H., 160 years old. Many large red maple were present in
the stand. The largest one, 13.5 inches D.B.H., was 70 years
old. Black ash ranked next in age being about 45 years old.
Reproduction of both black and green ash was abundant.
Red maple, northern white cedar, and balsam fir were also
doing well.

Plants of tho herbaceous layer were:

12221
Abundant --- Agg§_ggpggm (341)
Common ----- Frgxinug nigg; (2)
Rare ----~~- Apia; balsamea (9) Ulmus thgmasii (22)

 

Betgia‘ifiigngi) _

Sbrubs and Vines

Frequent --- Parthenocissus inserts
Rare ------- Amelanchier intermedia Corylus cognuta

Nemoganthus muoronata Rhus radicans
Ribes lacustre

 

Pteridonhytes
Common ----- Dgyopteris spinulosa
Rare ------- Botrychium dissectum DryOpteris thelypteris

 

 

Onoclea sensibilis

Sedseg. Grasses, and Herb;

Abundant --- Lysimachig terrestrig Maianthemum canadensis

Viola blanda

Frequent --- Carex intumescens Cornus canadensis
Fragarig vigginiana
Common --—-— Agrostis hyemalis Carex trisperma

Coptis groenlandica Galium obtgsum
Glyceria pallida

Rare ------- Solidago ulmifolia
c.Ash-Elm-Basswood stand
Location ----- Roscommon County, Lake Township,
T23N, RAW, SE 1/4, Section 33.
Soil type ----- Bergland loam.

Predominant species in this quadrat were the green

and black ash, slippery and American elm, and basswood. Few
birchos (Betula 1233; and Betula lgntg) and northern white
cedar, one small red maple completed the stand composition.
Three or four medium sized hemlock were dead and remained

standing.

The largest trees were found to be mostly ash and

elm, both approximately 60 years old.

Plants in the four milacre wuadrats were listed as:

... 'V‘

Frequent -—-

Common -----

Abundant -—-

Frequent —--

Common -----

Abundant ---
Frequent ---

Common -----

Abundant ---

Frequent —--

Common —----

Careximiusssssns

6O

Trees
Fraxinug pennsylvanica (l4)
Fraxinus nigrg (10)

Acer rubrum (l)
Betula lute; (2)

Shrubs and Vine;

_ ~q..lona

Parthenocissus inserta y
Rubus pubesceng .

Ribes americanum

Cornug stolonifera
Ijex verticillata

massages
Pteridophytes
Dryopteris thelypteris
Onoclea sensibilis
Eggisetgm sylvaticum
Sedges, Grasses, and Herbs

[mpgtiens pallidg

I Lycepus rubellus
Lysimachia thyrsiflorg

 

Calamagrostis canadensis Ehpatorium purpureum

Galium obtusum Glyceria pallida
Viol; blandg

Bidens cernua
Carex trisperma
Fragaria Virginian; var. illinoensis
Lysimachig nummularia {Lemma minor

Agrostis hyemalis

 

Scutellarig lateriflora Lemna Erasulcd. it
Solidago ulmifolia Spircdela polyrhiza

Agter lateriflorus Maianthemum canadensis

Sium suavg

- -
u - a
— a
... ,
a -
H 1

 

61
d.Elm-Maple stand
Location ----- Roscommon County, Lake Township,
T23N, R4W, SW 1/4, Section 13.
Soil type ----- Bergland clay loam.

Red maple, silver maple (Acer saccharinum),and slippery
elm were the dominant species with both black and green ash
as associate species. Two American elu. one balsam fir com-
pleted the tree species of the quadrat. A few Carpinus
caroliniana and one Amelanchier canadensis were the only
occurring shrubs.

The largest tree was a red maple, 26 inches D.B.H.,
about 90 years old. A 19.9 inches D.B.H. silver maple was
approximately of the same age. A 10.8 inches green ash, the
largest among the ashes, was about 60 years old. The American
elm was the oldest in the stand at 100 years.

Vegetation of the herbaceous layer were:

2222s

Abundant --- Acer rubrum (962) Ulmus sp. (451)
Fgaxinus nigrg (35)

Common ----- Fraxinus pennsylvanica (3)
Shrubs and Vines
Rare ------- Smilax tamnoides var. hispida
Pteridophxteg

Abundant --- gnoclea igngipilig
Common ----- Osmunda regalis

 

 

——~

 

62

Sedges,gGrasses, and Herbs

 

Carex intumescens Lemng trisulca

Cicuta bulbifera 'Spirodela polyrhiza
Lycogns americanus

Abundant --- Bidens frondosa iLemna minor

 

 

 

Frequent --- Agrostis hyemalis Galium obtusum
Impatiens capensis Lysgmachia thyrsiflora

Common ----- Geum aleppicum var. strictgm

Rare ------- Coptis greenlandica Glyceria striata

 

 

Maianthemum canadensis, Sium suave
Viola blanda

88

——

—
:hn‘ _
.l’:' .es

—

ties balsa

k5? Nbrum 3

3” 011nm
F"Hr‘nl'l‘lls n1 .
p'pemsYlv -
or“ F
awe‘ maria:
:1”
“(qua Decid' ‘

9h

“‘1‘ “EM

 

Table 2. Summarization data of four 1/5 acre Swamp Hardwood stands by size classes.

 

 

 

 

 

 

 

 

 

Size Classgg Totals
47 ' Impor-

§pecies greg. Dgnséiv §reg° Dingéiy g TFrgguqugr TSDgpgity Dr Ft6§§§21.A8:7acre Vgigge
Abies balsamea 3 75 17 11.4 .2 50 42 10.0 2 50 20 6.1 3 75 7-9 79 19.8 8.2 4.5 3.2 5.6 19.3
Acer rubrum 3 75 14 9.4 3 75 34 8.1 3 75 29 8.8 2 50 26 41.3 4 100 10.5 103 25.8 10.7 36.2 26.0 45.2 47.2
Acer saccharinum 1 25 2 1.4 1 25 2 0.5 l 25 1 0.3 1 25 5 9-5 1 25 2.7 11 2.7 1.2 10.2 7-3 12-8 11-2
Alnus rugosa 1 25 2 1.4 l 25 3 0.7 l 25 2.6 5 1.2 0.5 ~—-- -—- ———- 3.1
Amelanchier'sp. 1 25 4 2.7 2 50 6 ‘1.4 2 50 5.3 10 2.5 1.0 0.1 0.1 0.1 6.4
Betula lenta 1 25 l 0.2 1 25 4 1.2 1 25 2.6 5 1.2 0.5 0.8 0.6 1.0 3.7
giggiguiutea 1 25 3 2.0 1 25 3 0.7 2 50 13 3.9 2 50 5.3 19 4.7 2.0 3.0 2.2 3.7 9-5

caroliniana 1 25 8 1.9 1 25 2.6 8 2.0 0.8 0.3 0.2 0.3 3.6
Corylus cornuta 1 25 8 5.4 1 25 2.6 8 2.0 0.8 —-- --— --— 3.4
Fraxinus nigra 3 75 53 35-8 3 75 145 34.8 4 100 81 24.5 3 75 5 7.9 4 100 10.5 284 71.0 29.6 19.9 14.3 24.9 54.4
F. pennsylvanica 2 50 24 12.6 3 75 58 13.8 4 100 58 17.5 2 50' 5 7.9 4 100 10.5 145 36.2 15.1 17.7 12.7 22.2 38.3
Picea mariana 1 25 1 0.3 4 1 25 2.6 1 0.2 0.1 0.1 0.1 0.1 2.8
Thuja occidentalis l 25 16 10.8 3 75 71 17.0 3 75 64 19.3 1 251 3 4.8 3 75 7.9 154 38-5 16.0 14.4 10.3 18.1 34-2
Tilia americana l 25 2 1.4 2 5O 14 3.3 2 50 14 4.2 1 25 1 1.6 2 50 5.4 31 7.8 3.3 3.7 2.7 4.6 11.4
Tsuga canadensis l 25 1 0.2 1 25 4 1.2 1 25 V 2.6 5 1.2 0.5 0.5 0.4 0.6 3.5
Ulmus americana 1 25 1 0.7 3 75 3 0.7 2 50 4 1.2 3 75 7 11.1 4 100 10.5 15 3.8 1.6 11.6 8.3 14.4 20.4
Elgnggrubra 1 25, 2 1.4 3 75 28 6.7 3 75 38 11.5 3 75 10 15.9 3 75 7.9 78 19.5 58.1 16.2 20.3 20.3 27.6
____1 — —- 148 15.4 - -- 419 43.6 — -- 331 34.4 - 7’ 63 6.6 - 950 100.0 961 ‘--- 100.0 139.2 100.0 173.9 300.0

 

 

 

II-'|‘

 

64

Table 3. aummary data of the vegetation in milacre quadrats of thetfiump ng§vood stands as recorded in percentage of coverage or number of stems by height classes.

 

 

 

 

 

 

 

 

Total Black Ash-Wbite Cedar—Balsam Fir Ash—.Red M. 1e- Stand Ash-ElmBasswood Stand male M
Freq. 1 2 5 214 5 7 8 9 10 11 12 13 11. 15 16
Species (5) 0-1 1-3 0-1 1-3 0-1 1-3 0-1 1-3 0-1 1—3 0-1 1-3 0-1 1-3 0-1 1-3 0—1 l~3 0.1 1—3 0-1 1-3 0-1 1-3 0—1 1-3 0-1 1-3 0-1 1—3 0—1 1-3
Reg; 1
Abies balsamea 31.2 6 6 2 h 1 9
Acer rubrum 81.2 109 76 52 62 68 75 68 130 1 172 221 317 252
Betula lutea 31.2 5 3 2 1 2
Fraxinus nigra 68.8 7 6 A 1 1 1 2 3 4 1 10 2 8 9 5 1
Fraxinus pennsylvanica 43.8 5 7 5 2 6 1 1 2
Quercus rubra 6.2 1
Thuja occidentalis 18.7 1 8 1
Ulmus rubra 18.7 1 l 1
Ulmus thomlsii 31.2 ’22 1&0 126 2 112 A 67
Shrubs & Vines
W 18.7 3 3 1 T 1
Amelanchier intermedia 6.2 ‘ 22
Clematis Virginians 12.5 T T t
Cornus stolonifera 12.5 ' 1 l 2
Corylus cornuta 6.2 4
flex verticillata 25.0 1 1 3 2 5
Lonicera canadensis 6.2 1
Mitchella repels 12.5 2% T
Nemopanthus mucronata 6.2 1
Parthenocissus inserts 50.0 3% 3% 2% 6% 2% 2% T T 10%

Rhus radicans 12.5 ‘

 

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____‘ _=—‘_“ #3
Total Black Ash-White Cedar-Balsam Fir Ash-Red Ms 1e—E1m
___________.______.._________._...
2

Stand Ash-ElmrBasswood

 

 

Stand Elm-MB ls

 

65

‘ St d

 

 

32.3.1” Fag“ 0_1 3’3 '04 1-3 0-131-3 04414 02151-3 04 1__‘3 15933—3 0-1 1-3 0-1'1-3 0-1 1-3 0—1 1-3 0—1 1—3 04 143 0.1 1—3 03—1 1—3
Ribes americanul 18.7 2 5

Ribes lacustre 18.7 T T T

Rubus ideaus 6.2 T

Rubus pubescens 50.0 T 5% 5% T 8% 2%

Smilax tamnoides vamhispidug.2 T

Pteridophzges

Botrychium dissectum 12.5 T T

Cystopteris bulbifera 6.2 5%

Dryopteris cristats 12.5 T T

Dryopteris spinulosa 25.0 5% 3% 3% 25%

Dryopteris thelypteris 31.2 15% T 3% 10%

Equisetum sylvaticum 12.5 10%

Onoclea sensibilis 62.5 3% 2% 10% 10% 5% 20% 5% 3% 3% 1% T \
Osmunda regalis 12.5 35% 105‘
-S—=-—-—-———i;:.::.f:;:;:i.i .2. 1. 2,. 5. 1. 1. . 2% 1% 3“
Calamagrostis cahadensis 18.7 T 1% 5%

Carex intumsscens 81.3 2% 8% 1% 3% 1% 3% 3% T 3% 15% 1% 5%
Carex trisperma 25.0 T T 3%

Glycerin pallida 31.2 10% 2% 15% T 1% T

Glycerin an... 18.7 1% 3% 1% T

Aralia nudicaulis 12.5 5% 10%

Aster lateriflorus 25.0 1% 2% 1% T T

Aster puniceus 6.2 T

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Bidens cemua

Lemna minor
Lema trisulca

Mitella nuda

Rannueulus sp.

To.“ Ash-Red 1.— maasawood Stand Elm-Ma. 21 e Stand
Frog. 2 3 7 8 9 1o 11 12 13 11. 15 '13“
(1) 9-1 1-3 0-1 1-3 0-1 1.3 0-1 1.3 0—1 1-3 0-1 1-3 0—1 1-3 0-1 1-3 0-1 1-3 0-1 1—3 0-1 1-3 0—1 1—3 0-1 1-3 0-1 1.3 0-1 1.3 0-1 1—3
12.5 7! T 1% T
Bidens frondosa 25.0 7 1% T 1% 1% 1%
Cicuta bulbifera 25.0 T T T 1%
Coptis groenlandica 18.7 2% T 5%
Comus canadensis 37.5 3% T T 2% 3% T 2%
Eupatorimn purpureun 18.7 ‘ T 2% T 2%
Fragaria Virginians. 18.7 T 1% T T
12'5 T T
F. Virginians var. illinoensis
Galium obtusum 50.0 T 1% T T T T T T
Galium triflorun 25.0 2% T 1% T
Galium trifidun 18.7 '1' 1% 1%
Geun slappiun .m..strictu312.5 T T
Impatiens capensis 18.7 T T T
Impatiens pallida 25.0 3% 1% 3% 15‘ 25‘ 1% 2% \\
37.5 5% 1% T T T '1'
Spirodela polyrhiza
Lycopus emeticanus 75.0 2% 5% 5% 1% 1% 2% T T 3% 3% 2% 2% 1%
Lycopua rubellus 25.0 5% 2% 3% 1% 3% 1% 5% T
Lflimchia mammal-1a 12.5 1% T
Lysimchia thyrsifipra 68.8 T T T T 1% 3% 1% T 1% 1% 1% 1% 1% T T 1%
Msianthemm canadensis 56.2 T T T T 1%. 1 T T T T
18.7 T 10% 1%
6.2 T
Scutellaria laterifolia 13.7 T 1 5% T

J

 

 

 

 

 

 

 

 

 

 

67
Total Black Ash-White Cedar-Balsam Fir Ash-Red Ma le- Stand Ash—Elm*Basswood Stand Elm-Maple Stand
Freq. 1 2 3 1. 5 —7_—T" W 13 11. 15 T
Sgecies (%) 0-1 1—3 0—1 1-3 0—1 1—3 0—1 1-3 0—1 1-3 0—1 1-3 0-1 1-3 0-1 1-3 0-1 1-3 0-1 1—3 0—1 1-3 0-1 1-3 '0-1 1-3 0-1 1-3 0-1 1—3 0—1 1-3
Sium suave 12.5 I T T
Solidago ulmifolia 25.0 T 1% I 1% T
Thalictrum dasycarpum 6.2 1% T
Viola blanda 62.5 1% 2% 1% 1% . 1% T 1% 2% T T
T --- Denotes "Trace" which indicates less than 1% Coverage. ‘
1
1
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...-o‘

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68

2.8wamp Conifers of Black Spruce-Balsam Fir-Northern White
Cedar Association

Dominant species in this type were the balsam fir,
northern white cedar, tamarack. and black spruce. Red maple
and yellow birch.were the associate hardwoods. Four quadrats
of this type were studied:

a.Tamarack stand
Location ----- Crawford County, Beaver Creek Township,
T25N, R3W, SE i, Section 29.
Soil type ----- Rifle peat. T

Tamarack was the only tree species. The average age
of the stand was approximately 30 years old, with the largest
individual measured 9.6 inches D.B.H., and the smallest 1.3
.inches D.B.H.. One important fact was that no tamarack re-
production has been observed in the quadrat.

There was a dense shrub growth of green alder (Alnus
sinuata) and red osier dogwood (Cornus stolonifera). Some
swamp birch (Betula pumila) and Labrador tea (ledum greenish-
giggm) were also present. Sphagnum moss (Sphagnum sp.) was
abundant with a coverage of approximate 40 percent.

Soils were very wet and spongy with most of the herba-
ceous plants grown on the hummucks. Plants in the milacre
quadrats were: A

Irees
Rare ------- Quercus rubra (l)

 

—~

 

69

Shrubs and Vines

Abundant --- Rubus pubescens

 

 

 

 

Frequent --- Betula pdmila grunus Virginians
Logicera villosa var. solonis

Rare ------- Alnus sinuata Cornus stolonifera
ledum groenlangicumj i Prunus‘avium _

 

Spiraea latifolia
Pteridophytes
Common ----- DEXOpteris cristata

Sedges,_Grasses, and Herbs

 

 

 

 

 

Abundant --- égrostis hyemalis Bromus ciliatus
Calamagrostis canadensis Carex intumeggens
Galium aparine Maianthemum canadensis
Solidago uliginoga Solidago ulmifolia

 

Frequent —-- Campanula aparinoides

Rare ------4 Coptis greenlandica 'Lycopug americanus
Mitella nuda

b.Balsam Fir-Tamarack stand
Location ----- Crawford County, Beaver Creek Township,
T25N, R3W, SW 1, Section 35.
Soil type ----- Newton loamy sand (approximately 8 inches
layer of organic soil on top).
This stand was composed mostly of smaller trees.
Balsam fir was the predominant Species with the largest one,
8.4 inches D.B.H., about 40 years old. The largest of the five
tamaracks was 917 inches D.B.H., only 25 years old. The only
northern white cedar, 7.8 inches D.B.H., was 36 years of age.
Although there were nine different species present in the

quadrat, six of them were represented by a single tree.

 

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70

The predominant shrub species was the speckled alder
(Algal gugosa). Fire cherry (Prungs pensylvanica) and choke
cherry (grunge Virginians) were also abundant. These formed
a very dense growth.
large number of plants were present in the four mil-
acre quadrats. They were:
2:221
Abundant -—- Abigg balsamea (27)
Frequent --- Acer gubrum (15) Ulmus Sp. (6)

Rare ------- Betula lutea (ll) Quercus rubra (2)
Thuja occidentalis (2)

Shrugs and Vines

Abundant --- Cornus alternifolia Ilex verticillate
IThnnsflvigginiana-;fiN‘-Rnbugfpgbescens

Frequent~-- Alnus rugosa
Rubus ideagg var, acglegtigsimus

Common ----— Goran; stolonifega Lonicera villosa var. solonig
Nemopanthus mucgenata Ribes lacustre

Rare ------- Ribes americgggm

Ptegidophytes
Abundant -—- Dryopteris cristata var. clintoniana

 

 

 

Rare ------- Drygpteris spinulesa
SedgesI Grassesll and Herbs

Abundant --— Agrostis hyemalis Bromus ciliatus
Calamagrostis canadensis
Carex intumescens Fragaria vegca
ggligg trifloggm Glycegia boggalis
Glyceria striata Lycopus americanus

Maianthemum canadensis Mulenbergia racemosa

 

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Common ----- Aster sp. Geum aleppioum var. strictum
Impatiens capensis Lysimachia terrestris
Mitella nuda Ranuuculus sp.
Solidago ulmifolia Vina blanda
Rare ------- Aralia nudicaulis Coptis greenlandica
Goodyear pubescens Iris versicolor

 

 

Scutellaria lateriflora Solidago canadensis
Solidggo patula

c.Black Spruce-Balsam Fir stand
Location ----- Crawford County, Beaver Creek Township,
T25N, 34w, SE 1, Section 34.
Soil type ----- Rifle peat. -

This was a well stocked stand in which black spruce
and balsam fir were dominants. Some northern white cedar, a
few red maple and yellow birch completed the tree species of
the stand. The largest tree was a northern white cedar, 15.8
inches D.B.H., and the largest black spruce was 9.6 inches
D.B.H..

The stand was composed of 85 years old northern white
cedar, 50 years old balsam fir, and 40 years old black spruce.
The balsam fir had far more reproduction than the others.

Some seedlings of black Spruce were also present.

A few speckled alder were the only shrub species which

located at an opening in the stand. Vegetation of the milacre

quadrats were:

Trees
Abundant --- Abies balsamea (#0) Ace; rubrum (341)

Quegc gs rubra W(10)
Rare ------- Pinus Strobus (1)

 

 

 

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72

Shrubs and Vines
Abundant --- Vaccinium myrtilloides
Frequent -—- Rubus pubescens

Common ----- Ilex verticillata Linnaea borealis var.americana
Mitchella repens

Rare ------- Gautheria procumbens Lonicera villosa var. solonis
Prunus virginiana Ribes trieste

Pteridgphytes

Frequent --- gguisetum sylvaticum

Rare ------- Dryopterig cristata Dryopteris.§pinu19sa
Dryepteris thelypteris

Sedges. Gragges, and Herbs

 

 

 

 

 

 

 

 

Abundant -—- Aralia nudicaulis Carex intgmescens
Clintonia borealis Ceptis groenlandica
Cornus canadensis Glyceria borealis
Glyceria striata Maianthemum canadengig
gyrola seounda Trientalis borealis
Frequent --- Aster sp. Habenaria obtusata
Viola blanda
Common -—--— Mitella nuda Solidago uliginosa
Rare ------- Galium asprellum Geum aleppicum var. strictum

 

 

d.Northern white Cedar-Balsam Fir stand
Location ----- Roscommon County, Gerrish Township,
T24N,'R3W, NW &, Section 2.
Soil type ----- Rifle peat.
This stand was densely stocked with northern white
cedar and balsam fir as the dominant species, which associated

with black spruce and yellow birch.

 

 

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The largest tree was a 10 inches D.B.H. northern
white cedar. The stand was uneven—aged, consisting of 65
years old northern white cedar, 50 years old balsam fir, and
40 years old black Spruce. Both northern white cedar and
balsam fir were well represented in different diameter classes
and were well reproduced. Seven black Spruce were tallied in
the quadrat, all of them fell into size class 4 with no re—
production to be found. A few red maple and cherries were
scattered in the stand to complete the composition of tree
species.

A considerable number of Speckled alder was the sole
representative of shrub Species.

Plants in the four milacre quadrats were:

 

Trees
Abundant --- Abies balsamea (47) Acer rubrum (35)
Rare ------- Quercus rubra (l)

 

Shrubs and Vines

Frequent --- Lonicera canadensis Fubus pubescens

 

Common ----- Linnaea borealis var. americana
Rhus radicans

 

Rare ------- Acer spicatum Corylus cornuta
Ilex verticallata Mitchella repens

Ribes lacustre
Pteridophytes
Frequent --- Dryopteris thelypteris

Rare ------- Dryopteris Spinulosa

 

1)

 

 

74

Sedpes, Grasses, and Herbs

Abundant --- Acrostis hyemalis
Carex intumescens
Galium triflorum
Mitella nuda

 

Frequent --- Coptis groenlandica

Cemmon ----- Cornus canadensis
Prunella vulgaris
Viola blanda

Aralia nudicaulis
Clintonia borealis
Glyceria borealis

Eyrola secunda

Maianthemgm canadensis
Trientalis borealis

 

 

Table 4. Summarization data of four 1/5 acre Swamp Conifers stands

by Size classes.

 

 

 

 

 

 

 

 

SIZE CLASSES TOTALS

2 3 4 5h m Impor—

5mm fl W 1.55% %—-%1- §——% Dan-$61- .§-“% 19%?” Triliuenéif T D9331” Tm a~§§-%_;T%‘%%%7x viii?
Abies balsamea 5 75 42 75.7 5 75 301 60.0 5 75 94 24.5 5 75 15.6 437 109.2 45.9 21.5 26.6 26.6 86.1
Acer rubrum 1 25 3 5.3 l 25 1 0.2 2 50 15 3.4 1 25 1 12.5 3 75 15.6 18 4.5 1.9 3.1 3.9 3.9 19.4
Betula lutea 1 25 5 1.0 2 50 11 2.8 1 25 2 25.0 5 75 15.6 18 4.5 1.9 3.1 3.9 3.9 19.4
Fraxinus nigra 1 25 1 0.3 1 25 4.6 1 1.2 0.1 0.1 0.1 0.1 4.8
Larix laricina 2 50 45 9.0 2 50 100 26.0 2 50 2 25.0 2 50 9.1 147 36.8 15.5 19.0 23.7 23.7 48.3
Plcea mariana 1 25 2 3.5 1 25 20 4.0 5 75 41 10.7 1 25 1 12.5 3 75 13.6 64 16.0 6.7 9.7 12.1 12.1 32.4
Pinus Strobus 1 25 l 0.33 l 25 4.6 l 1.2 0.1 0.1 0.1 0.1 4.8
Prunus serotina 2 50 7 12.2 2 50 8 1.5 2 50 9.1 15 3.8 1.6 0.1 0.1 0.1 10.8
Thuja occidentalis 1 25 5 5.3 2 50 122 24.3 3 75 122 31.7"2 50 2 25-0 3 75 13.6 249 62.2 25.2 23.3 29.1 29.1 68.9
Ulmus americana l 25 l 0.1 l 25 4-5 l 1.2 0.1 0.3 0.4 0.4 5.1
_ __ 57 6.0 _ __ 502 52.8 - _- Egg 40,4 - —- 8 0.8 —550 100.0 951 --—- 100.0 80.1 100.0 100.0 500.0

 

75

76

 

 

 

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8H was. 0.3 3. ads 3 E. «a a 6.8 3 mm a n.» mm 3 a 33332. 395
R .3 .2. «.m fin 3 4.3 R a flu .. on N «J. 2 an a 3331.8 88cm
3 «J Sm o; m; a. 4.3 R a n: a. R a £3333. 558
a v.0 a; do m6 a E. 3 a «.o a «a a .... 2223...:
3a «3“ 0.2. «.9 «.8 co 5. mm a 5.3 cu mu a 93 2. «a a 335. 2:5
2.3 mg. 0.2 0.2. «.93 an 5.8 8H 4 «.3. can 03 a «J.» 03 2.: 1. Some» use:
a «A o.~ m6 To a 5. mm a ...o 3 a seasons .31
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Table 6. Sum

3931"

g

'22:

7'; balsamea

:2.- mm

’11! lutea

I533 Strobus

"9”“ rubra

liji "adultelis

7’4 thou.“

W

or 'Picstu.
Q“ "goes
2“?“ 'hmta

 

 

T‘ble 6.

 

 

77

Summary data of the vegetation in milacre quadrats of the Swamp Conifer stands as recorded in percentage of coverage or number of stems by height classes.

 

 

 

 

Total Tamarack Stggg Balsam Fir-Temarack 7 sgand Blagk Sgruceigalsam Fii Signd Whit; Cedarelesam Fi§5 Siand

§£ecies Fig? 0411.3 0-1 1-3 0431-3 0411.3 0-151-3 0-1 1—3 0-1 1—3 0-1 1—3 o~1 1—3 0—1 1.3 0-1 1.3 0—1 1-3 0—1 1—3 0-1 1—3 0-1 1—3 0~1 1—3
ggffi balsamea 75.0 1 l 11 1h 10 18 8 h 5 10 16 . 16
Acer rubrum 68.8 11 2 l 1 90 128 Al 82 10 9 l2 1 3
Betula lutea 6.2 11
Pinus Strobus 6.2 l
Quercus rubra h3a8 l 2 5 l 3 l 1
Thuja occidentalis 6.2 2
Ulmus thomasii 18.7 3 1 1
samba & Vines 1 ‘
Acer spicatum 6.2
Alnus rugosa. 18.7 7 1 2 1
Alnus sinuata 6¢2 l
Betula pumila 18.7 7 l h s“~_‘_“‘
Cornus alternifolia 25.0 15 21 20 5 3 3 14 4
Cornus stolonifera 18.7 3 1 5 1
Corylus cornuta 6.2 1
Gautheria procumbens 6.2 T
Ilex verticillata h3o8 2 3 1 2 l A l
Ledum groenlandicum 6.2 2
Linnaea borealis T T T T

var. americana 25.0
Lonicera canadensis 18o7 1 3 1 1
L. villosa var. solonis 37.5 2 1 1 2 1 1
Mitchella repene 18.7 1% T T

 

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78

 

 

 

 

Prunus avium

Rhus radicans

Ribes lacustre

Ribes trieste
Rubus ideaus

Pteridophztes

Herbs

Aster sp.

Fragaria vesca
Galium aparine

Galium asprellum

Total Tamerack Stand Balsam F535EEEEFack Stand Black.SEruce-Balsam Fir Stand N. fiiite CedarnBalsam Fir Stand
Freq. ‘3“ 2 3 T 7“"?- 1 e 9 10 11 12 13 u, 15 16
(2:) 0—1 1—3 0-1 1—3 0-1 1—3 0-1 1—3 0—1 1-3 0—1 1-3‘ 0-1 1-1 0-1 1-3 0—1 1-3 0~1 L3 0-1 1—3 0—1 1—3 0—1 1—3 0-1 1—3 0-1 1—3 0-1 1—3
Nemapanthus mueronata 12.5 1 3 1 1
6.2 2
Prunus virginiana 50.0 2 l 3 3 5 l l 1 1
12.5 3% T
Ribes americanum 6.2 1%
18.7 1 1 l
6.2 1
var. aculeatissimus 18.7 3% 5% 10% 5%
Rubus pubescens 87.5 15% 10% 20% 15% 10% 5% 10% 10% 15% 15% 5% 15% T 40% 2% 30% T 3%
Spiraea latifolia 6.2 1
Vaccinium mtilloides 25.0 25% 1% 1% T 2%
Dryopteris cristata 18.7 T T 1%
”12518333323? 25.0 1% 1% 3% 1%
Dryopteris spinulosa 18.7 T 1% 1% ‘~——
Dryopteris thelypteris 25.0 5% 2% T 2%
Equisetum sylvaticum 18.7 5% T T
ma nudicaulis 56,2 1 T 10% 5% 5% 10% :51. 5% 5% 31% 31% 5% 152% 5% 8%
31.2 T 1 T T T 1%
Aster junciformis 18.7 T T T
Clintonia borealis 50.0 l 1% 2% 2% T 3% 3% 3% 3%
Coptis groenlandica 56.2 T 3% L 2% 1% 3% 1% 5% T 1%
Cornus cenadensis 37.5 E1 T 1% T T 3% T T 1%
25.0 } 1% 3% 1% T
25.0' 10% 20% 15% 10% ,
6.2 i T

 

 

79

 

 

- ‘16; 01%

 

 

 

IF“ nun-

Galium triflorum
Geum aleppicum

Iris versicolor

Mitella nudl

Pyrola secunda

Rannuculus sp.

Solidago patull

Viola blanda

Sedges & Grasses

Bromus ciliatus

Glyceria strilts

Iotal EggggggL Stggg 001eez;g£¢Teeareek Stcnd Black Sgruce~Balsam Fir Stand N fiilte Cedareaaleam Fir Eiiii
?ren. 1 2 ‘3 1 “”'?‘"fl '7§‘""""7“""“§“‘ 3.._. 10 11 12 ’a"i5'""'"1Zf’—""15_-‘_'_'ig—'
(£J 0~1 1—3 0~1 1.3 0~1 1_3 ~~1 1—3 ,-1 T-j 5-1 Tug. 0-1 1—3 0-1 1.3 0—1 1—3 0-1 1~3 0-1 1-3 0-1 1.3 0-1 1~3 0-1 1—3 0—1 1.3 0-1 mes
50.0 1% T 2% 1% 3% 1% T T
var. strictum 18.7 T T T
Goodyear pubescens 6.2 T
Habeneria obtusata 18.7 T 3% T
Impatiens capensis 12.5 T 1%
6.2 1%
Lycopus americanus 31.2 10% 3% 1% 25% 2% 15% 2% 2%
Lysimachia terrestris 12.5 T T
Maianthemum canadensis 100.0 1% 1% 2% 1% T T T T 1% 1% T 1% 2% T T 1%
56.2 1% 5% L T 1% T 10% 2% 5% 2%
Prunella vulgaris 12.5 r 3% 3%
“3.8 T T T T 1% T T ~
12.5 T T
Scutelllris lateriflora 6.2 T
Solidago canadensis 6.2 1 T 1% “‘“—
6.2 T
Solidago uliginosa h3.8 T T T T T 1 T 1% T T
Solidago ulmifolia 37.5 T 2% 2% 2% 2% 2% T T T 2% 2%
Trientalis borealis 50.0 T 1% ' 2% 3% 2% T T 1%
13.8 T A T 2% T T T 2% T
W. W 28:8 éiéééfi/Wé//¢¢¢¢¢¢¢//////é}W?éWéf?W???éflfflfifiéfiéfiflffi
Agrostis hyemnlis 75,0 20% 30% 35% 10% ///// /////////////// / ///////// O - %
3331-1313322??? 13223 /////////// /////////////////////// 25" 50% W 2% 1% 1% 2% T 3% 27 3
32:8 éfiéfiéfiéfléfléfiféfifiéflfléfifléfl //////////////////////////////////MW555?fléfiéfifiéflffléfié/fi

Muhlenbergis rscemosa

 

T --— Denotes "Trace" which indicates less then 11 coverage.

115.35.... .. _ -

80

3.Lowland Aspen of Pepulus-figlig Association

Trembling aspen (Populus tremuloides) was the major
species in the type, with figlin élgug, and Cornus as the
dominant shrub species. Two stands were selected for quadrat
study. On the region as a whole, this type does not occupy
extensive area but occurs in small tracts here and there
bordering marshes or brooksides.

a.Trembling Aspen-Longebeaked Willow stand
location ----- Roscommon County, Lake Township,
T23N, R4W, SW i, Section 16.
Soil Type ----- Newton sand. ‘

This was a poorly—stocked trembling aspen stand of
approximately 30 years old. One birch was the only other tree
species found in the quadrat.

Alggg ruaosa and gall; bebbiana were the dominant
shrubs. A few red osier dogwood were also present.

Plants in the four milacre quadrats were:

‘ 11:929.

Abundant --- Betula lutea (21) Acer rubgum (l)
Quercus rubra (l)

Shrubs and Vines

Abundant --- Cornus stolonifera gig; verticillata
Eggug pubescens

Frequent --- glad; rugosa ‘figlig bebbiana

Common ----- 512g; lacustre

Rare ------- Lonicera oblongifolia Rhus radicans

 

 

 

Rubus ideaus var. aculeatissimus
Vaccinium vacillans

"S

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81‘

 

 

Pteridophytes
Common --——— Eguisetun palustre ' Drxopteris apinulosa
Rare ------- Onoclea sensibilis

 

Badges. Grasseg. and Herbs

Abundant --- Carex diandra Glyceria borealis
Lysimachig terrestris licoous americanug

Frequent --- Campanula aparinOides Fr aria Virginians
Pyrola rotundifolia Vio a blanda

Common ----- Aster lateriflgrus Galium sp.
Mentha grvensis erola secunda

Solidaao ulmifolia

Rare ------ - Agrostis hxemalis

Epilobium glandulosum var. adenocaulon .~
Maianthemum canadensis Scutellaria epilobiifolia

Smilacina trifolia figlidazo caesia E.
b.Trembling Aspen-Heartleaf Willow stand
Location ~e--- Roscommon County, Markey Township,
T23N, R3W, SE &, Section 1.
Soil type ~---- Newton sand.

Trembling aspen was the only tree species found in
this quadrat. This was a better stand than the first one
with more trees and better growth. The largest one was 7.7 '
inches D.B.H., 35 years old.

The Speckled alder and the heart-leaf willow (gall;
griocephala) were the dominant shrub species. .

Components of the herbaceous layer were:

22222

Rare — ------ Populus tremuloides (l)

 

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82

Shrubs and Vines
Salix alba var. vitellina

Alnus rugosa Cornus stolonifera
Salix ggyzdaloideg

 

Asclepias incarnata
Eteridophytes
Drxopteris spinulosa

Sedaes. Grasses. and Herbs

 

 

Carex intumescens Lysimachia terrestris
LxcoEus americanus Scutellaria epilobiifolia

 

 

 

Calamagrostis canadensis
Galium trifidum

 

 

 

Agropyron repens Campanula aparinoideg
Cicuta bulbifera Epilobium leptophxllum
Junous canadensis Lysimachia thyrsiflora

 

 

Scirpus experinus

Aster juncigormig Eupatorium perfoliatum
Fragaria Virginians Rumex verticillatus

-..

 

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§ 83
W

Table 7. Summarization data of two 1/5 acre lowland Aspen stands by size classes.

 

 

 

 

 

 

 

sxzs CLASSES ' TOTALS

Impor-

Betula lutea 1 50 1 1.7 ; 1 50 33.3 1 0.5 1.2 0.120 1.2 0.30 35.7
Populus tremuloides 2 100 24 100 2 100 58 98.3 2 100 66.7 82 41.0 98.8 9.543 08.8 23.86 264.3
— --— 21+ 20.0 - ~—- 59 71.0 — 150 100.0 83 —_—- 100.0 9.663 100.0 24.16 300.0

 

Table 8. Summarizstion data of two l/IO acre lowland Aspen stands by height classes.

 

 

 

 

 

 

 

 

HEIGHT CLASSES 'M TALS
6—15 Feet 15-25 Feet Abun-
§pecies giggg; i%§2§l%%- A§£§%; _§g§§i%%_ TEE§%%3§%%§_ 53‘2§%§l£1656” dance Ar #/AOPe
Alnus rugosa 2 100 157 80.5 2 100 246 66.8 2 100 33.:3 403 201.5 71.6 201.5 56.6 2015
Cornus stolonifera 2 100 11 5.6 - -—— —-— —-—— 2 100 33.3 11 5.5 2.0 5.5 1-5 55
Salix bebbiana 1 5o 7 3.6 1 50 80 21.7 1 50 16t7 , 87‘ 43.5 15.4 87.0 24.5 435

I

Salix eriocephala 1 50 20 10.3 1 50 42 11.5 1 50 16.1 i 62 31.0 11.0 62.0 17.4 310
. 'I

- --- 195 100.0 - —-— 368 100.0 - 300 100.: 563 ———— 100.0 356.0 100.0 2815

 

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88

4.3wamp Shrubs of figlig-Cornus-Alnus Association
This type consisted a mixture of willow, dogwood,
alder, swamp birch, etc., to form a dense thicket growth.
Only one stand was selected for quadrat study. It
was bordering a tamarack swamp and along the stream. All
plants were below 6 feet in height, therefore only milacre
quadrats were being studied.
Location ----- Crawford County, Beaver Creek Township,
T25N, 33w, Nw i, Section 33.
Soil type ----- Rifle peat.

Plants in the four milacre quadrats were:

Trees
Rare ------- Betula lutea (6)

Shrubs and Vines

 

 

 

Abundant --- Alnus rugosa Betula pumila
Cornus stoloniiera Lonicera villosa var. solonis
Potentilla fructicosa Rubus hispidus
Sajix petiolaris §piraea alba

Frequent --- Aronia nigrg

Common ~--—- Ribes trieste

 

 

 

 

 

 

 

Rare ------- Cornus alternifolia
PteridODhyteg
Rare ------- Dryopteris cristata
Sedges, Grasaes, and Herbs
Abundant --- Azropyron repens Bromus purgens
Muhlenbergia racemosa Solidago canadensis
Pregnant --- Galium asprellum Solidago uliginosa

Solidago ulmifolia
Common ~--~- Aster juncoformis
Rare ------- Aster novae-aggliag Circium muticum

-~'_--

 

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Table 10. Summary data or the feur milacre quadrats in the Swan Shrubs
stand by height classes. Plants were recorded in percentage ef
severags er number of atoms.

 

W

 

 

 

 

 

 

 

 

Total 1 2 '3 A
Freq. HEIGHT CLASSES
Species (5) 0-1 1.3 1—6 0-1 1;“; j-é 0-1 1-3 3—6 o-1 1-13-6
(Trees)
Betula lutea 25 6
(Shrubs and Vines)
Alnue rugeea 100 3 2 ' 1 2 2 2 1
Arenia nigra 75 5% 10% 1% 5% 5%
Betula manila 100 10% 15% 1% 15% 5% 10% 2% 10%
Cornus alternifelia 25 5%
Cernus stelsnifera 100 5% 10% 1% 5% 10%
Leaicera villesa
var. selenis 100 2% 3% 5% 20% 2% 10% T 5%
' Petentilla fructicesa 100 15% 25% 20% 15%
Ribes lacustre 50 '1‘ T
Rubus mm... 100 5% 5% 10% 3%
Salix petielaris 100 1% 1% 5% 1% 5% 1% 1% 3%
Spiraea alba 100 5% 5% 15% 10% 5% 5% 5%
(Sedgee, Grasses 8: Herbs)
Aer-Hr» rom- 100 ~81; —~ ---- 5% --- -- 5% --- --- 10% --
Brem pursue 100 ~------ 8% ---- --- 5% --- --- 2% -- --- 10% ---
Glyéeria berealie 75 --- 5% --- ---- 5% -- --- 5% “-
lluhleabergis racemesa 100 -- 5% ----~- ---- 2% -- ---- 8% ~— --- 10% --
Aster Juseinerlis 50 T T
Aster aevae—aaglhe 25 1%
Circiul autism 25 1%
Galin- aeprellu 75 2% 1% 1%

- te be eentisued ..

 

‘-—-‘-

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Preq. HEIGHT ~__ CLASSES
sponge fi (29) A 0-1 1-31-6 0-1 1-3 3-6 0-1 :3 3-6 04 1-2 27-6
Selidage canadensis 100 T T T T
Selidage uliginesa 75 . T T T T
Selidage ulmifelis 75 T T 2% 1%

( Pteridephitee)
Dgzezgegis cristataw 25 T

T -- Denetes "Trace" which indicates less than 1% severage.

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5.The Marsh or Open-Meadow type
Davis (1906) has defined the "marsh" and "bog" as:

\

”The 3bogi is an area of wet, porous land, on

which the soil is made up principally of decayed
and decaying vegetable matter, so loosely conso-
lidated, and containing so much water, that the
surface shakes and trembles as one walks over it.
The vegetation upon the surface is characteristi—
cally either some species of moss or sedge, or
grass, or a combination of two or more of these
with shrubs and even smill trees.

A 'marsh' has a firm soil, it may be soft and
very wet, even submerged, and the vegetation upon
it is principally grass-like. Shrubs may occur
but are not infrequently form thickets"

Three stands were selected for sample plotsjin this
type. Two were bogs and one was the marsh. The bogs were
found on Greenwood peat; and the marsh was on Newton sand.

a.Scirpus- ypha stand
Location ----- Roscommon County, Lake Township,
T23N, R4w, NE &, Section 21.
Soil type ----- Greenwood peat.

Dominant species of this stand were the grassy vege—
tation consisting of Scirpus cyperinus, several species of
Carex, and Calamagrostis canadensis. Juncus canadensis,
Eleocharis obtusa, Glyceria canadensis were also present with
the former two found on very wet sites.

The water-table of the soil was very near the surface
to form an almost permanent wet condition, as indicated by
such partly submerged plants as : Scirpus cyperinug, Carex

iasiocarpa, Eleocharis obtusa, Juncus canadensis, Typha

{I}

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.
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...—-...—

...—..---

92

latifolia, £21; versicolor, Bidens cernua, Alisma triviale,
and gagittaria montevidensis. Davis (1906) has classified
such type as the "Rush Swamp'.

” The secondary species were Typha latifolia, 12;;
versicolor, Sagittaria montevidensis, Qiguta bulbifera,
§;3m_§g§1gJ Lysimachia terrestgig, Eidens cernua, Hypericum
boreale, Lycopus americanus, Rumex vertigiliatus, etc.,

One Species of willow (gall; sp.) was the only shrub
present in the stand.

The occurring plants in the four milacre quadrats

 

 

 

 

 

 

 

 

 

 

 

 

 

were:
Shrubs-and Vines

Common ----- Salix sp.
SedgesLGrassesl and Herbs

Abundant --- ggdens cernua Calamagrostis canadensis
Carex lasigcarpa Cicuta bulbifera
Galium trifidum Lysimachia terrestrig
Sagittarig montevidensis Scirpus cyperinus
Slum suave Iypha latifolia

Frequent —-- Carex crinita Carex comosa
Eleocharis obtusa Glyceria canadensis
Hypericum boreale Iris versicolor
Juncus canadensis Lycopus americanus

 

Rumex verticillatus
Rare ------- Alisma triviale Scutellarialepilobiifolia

 

b.0alamaerostis-Scirpus stand
Location ----- Roscommon County, lake Township,
T23N, RAW, SW %, Section 21.
Soil type ----- Greenwood peat.
This was an ecologically slight more advance stand
than the previous one, as indicated by the increase of
galamagrostis canadensis, the presence of Agrostis h emalis,

Aster, Solidaqo, and the absence of such partly submerged

 

species as glisma, Sagittaria, Typha, Eleocharig, etc..
Willow (Salix discqlgg) remained to be the only
occurring shrubs but was more frequent in presence and also
increased in its coverage.
vegetation in the four milacre quadrats were:
Shrubstand Vines
Frequent --- gall; discolor

Sedges,_Grasses, and Herbs

 

Abundant —-- Aarostis hyemalis Calamagrostis canadensis
Carex lasiocarpa Cicuta bulbifera

 

 

Lysimachia terrestris Scirpus cyperinus
veronica scutellaria

Frequent --- Galium.trifidum Hypericum‘boreale
Scutellaria epilobiifolia
Slum suave

 

 

 

 

 

Common ----- Carex tenera Epilobium leptophyllum
Eupatorium perfoliatum Lycopus americanus
Rare ------- Aster Junciformis Aster sp.
Bidens cernga Carex comosa
Galium tinctorium Solidago rugosa

 

Solidaqo uliginosa Rumex verticillatus

a
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...-—

.c.§gl;§-Calamagrostig stand
Location ----- Roscommon County, Lake Township,
T23N, R4W, NW %, Section 21.
Soil type ----- Newton loamy sand.
This was a more advanced stage of the marsh type.
Although the soil is somewhat different than the peat, but
a relatively thick layer of organic soil on top of the wet
sand makes it similar to the peat. Water-table was very near
or at the surface during most part of the growing season.
However it draped to as deep as 24 inches in a dry period.
The increase of willows in tumber of species and in
coverages indicated a drier condition than the preceeding
two stands. This invasion of shrubs into the wet marsh area
is slow and may last for a long period of time.

' §alig petiolaris and Salix gegissimg were the domi-
nant shrubs which formed dense bushes. The diameter of their
spreading crown ranged from less than one foot to as large
as six, seven feet or more. Most of the willows attained a
height from 3 to 5 feet, a few also reached 6 feet or more.
The following data shows the height and diameter of the crown

of shrubs over 6 feet tall in a l/lO acre quadrat:

 

Height Diameter of
Species £Ft.) Crown LFt.)
Salix petiolaris 7 6
6 6.5
5 7

Salix serissima 5.5 6

Vegetation in the four milacre quadrats were:

 

Trees
Common ----- Betula lutea (6)
Rare ------- Populus tremuloides (l4)

 

Shrubs and Vines

Common ----- Salix alba var. vitellina
Salix serissima

Rare ------- Alnus ruqosa Salix lucida
Salix petiolaris Spiraea alba

 

Rubus flagellaris

Sedges, Grasses, and Herbs

 

 

 

 

 

 

Abundant --- Agrostis hyemalis. égter Junciformis
Calamagrostis canadensis Carex lasiocarpa
Carex tenera Iris versicolor
Lyccpus amergganus Lysimachia terrestris

 

 

Scirpus cyperinus

Frequent --- Cicuta bulbifera
Solidaeo uliginosa

Common ----- Eupatorium perfoliatum Galium trifidum
Hyfiericum boreale

Rare ------- Campanula aparinoides Solidago graminifolia
Solidago missouriensis

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ECOLOGICAL FACTORS

Ecology is the study of organisms in relation to
their environment. Therefore, in making an ecological study
of forest succession, it is necessary to investigate the
environment in addition to plant communities.

The site factors which are essential for the ecolo-
gical description and characterization of associations are
best considered under three groups:(l)Climatic or atmospheric
factors, (2)Edaphic or soil factors, and.(3)Biotic factors
or the effects of the living environment.

These factors are responsible not only for the ecolo-

gical characteristics of the present vegetation but also for

successional changes of the plant communities.

A.Climatic factors

The climatic facters relate to atmospheric conditions
and include all factors influencing plant life which are
associated with the atmosphere. Four factors are being con-
sidered in this paper. They are: temperature, relative humi-
dity, precipitation, and evaporation. Climatological data

pertaining to the region are shown in Table l.

 

 

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l.Temperature

Meriam (1898) stated that air temperature is the most
important factor in fixing the limits beyond which particular
species and particular communities can not extend. Weaver and
Clements (1938) have stated that "Temperature is like water
in its action upon plants in that it has more or less to do
with nearly every function, but as a working condition and
not as a material."

Observations indicate that the various tree species
can live only at temperature of a certain range, which may be
termed their critical temperatures. However, within a rela-
tively small, climatically similar area, such as the region
of this study, local variations in temperature are very
little which can be disregarded in their'direct'éffect on ve-
getational changes.

Yet, vegetation tends to moderate the temperature.
This equalizing effect increases with number and height of
the layers (Braun-Blanquet, 1932). Therefore temperature
is being considered as an influencial factor to the evapo-
ration rates of different vegetation types in this study.

Average weekly maximum and minimum temperatures for
the period from July 20 to October 4, 1952, as recorded from
the Higgins Lake and Houghton Lake Weather Bureau Stations
are shown in Table 12. Graphical presentation of the data

is shown in Fig. 6.

 

101

Table 12. Weekly averages of temperature, relative
humidity, and accumulated precipitation during
the period from July 20 to October 4, 1952,
compiled from data obtained at Weather Bureau
Stations near the Higgins Lake area.

 

 

Temperature UF. R. H. %' Precipitation-Inches
Week Hi%§;:s Hoigggon Ave- Rose Cit Higgins Houghton Ave-
(Ended) Max. Min. Max. Min. rage jlupton __Lake iggge rage
July 26 82.6 61.1 82.7 61.7 72.0 51.4 1.63 2.79 2.21
Aug. 2 76.8 51.6 77.0 54.3 64.9 59.1 0.19 0.12 0.16‘
‘9 75.1 51.1 76.3 53.7 64.0 65.3 2.24 2.27 2.25
16 78.4 53.1 78.4 55.3 66.3 58.7 ' 0.12 0.06 0.09
23 73.8 48.8 75.4 53.0 62.8 51.4 0.42 0.04 0.23
30 83.0 51.8 84.7 53.7 68.3 62.1 0.00 0.00 0.00
Sept. 6 72.7 53.1 73.3 51.5 62.7 65.7 0.98 1.19 1.08
13 83.3 51.1 81.8 51.8 67.0 54.7 0.00 0.00 0.00
20 68.3 46.1 70.3 47.3 58.0 68.1 0.21 0.48 0.35
27 63.8 38.6 65.2 38.0 51.4 65.0 0.56 0.47 0.51
Oct. 4 62.4139.8 65.8 40.0 552.0 57.0 0.26 0.23 0.24

 

 

 

 

 

Temperature 0?.

102

90

 

70

soy

50 ‘ ,’ \

hoe \ \ ’

 

35 1 1 1 L 1 1 1 L 1 1

20—26 27-2 3-9 10-16 17-23 Zip-30 31-6 7-13 ELL—20 21—27 28-1.
July Aug. Sept. -. "Oct.

0

 

 

...—... Maximum temperature ----- Minimum temperature

Fig. 6. Weekly data of maximum and minimum temperatures ae recorded
at the Higgins Lake and Houghton Lake Weather Statione
during the period from July 20 to October 1., 1952.

.--

r“
5";

103

. 2.Relative humidity

The only available data of the relative humidity
obtained from the Weather Bureau Stations near the Higgins
Lake area was the one at Rose Lake (Lupton), Ogemaw County.
Percentages of relative humidity were transformed from wet I
and dry bulb readings. Weekly averages of the period from
July 20 to October 4, 1952 were derived from daily data and
are shown in Table 12.

Humidity is affected by temperature, wind, altitude,
exposure, cover, and soil moisture content (Weaver & Clements,
1938). High temperatures lower the relative humidity; wind
has a powerful effect upon humidity in that dry winds lower
the amount of air moisture and promote transpiration, whereas
the moist winds exert an opposite influence; exposure affects
humidity through the action of sun and wind; cover increases
humidity by reducing the influence of both temperature and
wind; and evaporation from the surface of moist soils increases
humidity.

Generally speaking, forested regions have high humi-
dities; while the humidity of open area or graBSIand‘is low.
Lowlands are more humid; and tablelands and mountains usually

less humid.

104
3.Precipitation

The water requirements of plants are chiefly met by
the precipitation of the water vapor of the air in the form
of rain, dew, or snow. The atmospheric precipitation is of
particular importance in its effect on vegetation, owing to
its relation to atmospheric humidity and soil moisture con-
tent. In general, fluctuations in soil moisture are direct-
ly traceable to variations in precipitation.

Toumey & Korstian (1948) stated that

"Variations in geographical distribution of

precipitation influence markedly the distribution
of forest. With the same amount of annual pre-
cipitation the character of a forest depends upon
whether the rainfall is in the growing season or
in the cold season; whether it is evenly distri-
buted or confined to a few months of the year.

In fact the distribution of precipitation through
out the year, especially when rainfall is not
heavy, may determine whether a forest is present
or a type of vegetation needing less water."

However, in this study, precipitation would not be
an important factor in influencing the soil moisture because -
of the fact that the soils of the swampy lands are almost
continuously wet throughout the year. The annual precipita-
tion of about 30 inches in this region could not possibly be
responsible alone; edaphic factors must also be considered.

Weekly precipitation data at the Higgins Lake and
Houghton Lake Weather Stations during the period from July
20 to October 4, 1952 are shown in Table 12. Graphical pre-

sentation of the data is shown in Fig. 7.

Precipitation - In Inches

 

 

 

 

JJl

 

20-26 21-2. 3-9 )0-16 ”-23 “-30 31-6

Jutg

Aug.

1.13 14-20 21-27 26-4
Sq». Oct.

Fig. 7. Weekly accumulated precipitation during the period from

July‘ZG to October 1., 1952.

106

4.Evaporation

Braun-Blanquet (1932) stated that "the evaporation
rate is the combined effect of humidity, temperature, wind,
atmospheric pressure, and radiant energy." Evaporation
markedly determine the efficiency of rainfall, especially
where the annual precipitation is less than 30 inches
(Weaver and Clements, 1938).

Table 13. Evaporation rate as recorded by week at

stations of 3 different vegetation layers during
the period from July 20 to October 4, 1952.

 

-_

 

 

 

 

Evaporation in C.C. of Water
Date of Salix-Cornus- Thuja-Abies
Measurement Marsh Stand _§1nus Stand Stand
July 26 65.5 49.3 43.1
Aug. 2 98.6 57.7 48.5
9 37.7 16.2 1.3
16 77.7 30.8 28.0
23 91.6 35.4 30.0
30 85.5 50.8 43.1
Sept. 6 54.7 32.3 27.8
13 93.3 61.6 44.7
20 74.3 40.0 35.
27 55.3 28.5 17.7
Oct. 4 93.6 72.8 47.8

 

evaporation rate at three stands of different vegetation

layers in this study.

Livingston atmometers were used to determine the

The three stands selected for the

107

 

100 ..

CD
0
l

O\
O

40-

Weekly Atmometer loss — c.c.'s

m.
0
J;

 

‘I

 

 

O A K A a A A 1 1 L
20-26 27-2 3-9 10-16 17-23 24-30 31-6 7-13 14-20 21-27 28-1.
July Aug. Sept. Oct.

 

Harsh type
Swamp Shrubs type

.1-

S.... Swamp Conifers type

 

Fig. 8". Weekly evaporation in cubic centimeters in Marsh type,
Swamp Shrubs type, and Swamp Conifers type for the
"peridd from July 20 to October 1., 1952.

1.51‘

.. ...-.-

108

experiment were: (l)The Marsh. (2)The Swamp Shrubs of Sgllg-
Cornus-Alnus stand. (3)The Thuja-Abies stand of the Swamp
Conifers.

Photographs of atmometer set up in the first two
stands are shown in Plate'lwand 2. All atmometers were set
up with bulbs about 18 inches above the ground surface.

Weekly readings were recorded during the period from
July 20 to October 4, 1952, as shown in Table 13. Graphical
presentation of the data is shown in Fig. .8.

Table 14. Analysis of variance of the evaporation

data obtained from three different stands during
the period from July 20 to October 4, 1952.

 

 

 

Source of Degree of Sum of Mean

Variation Freedom Sguares Sguare F
Total 32 19505.33 609.54

Between stands 2 10584.32 5292.16 lOO.l4**
Between weeks 10 7864.00 786.40 14.88**
Error 20 1057.01 52.85

 

**Denotes significant at 1% level.

Statistical analysis in Table 14 shows highly signi-
ficant differences between the evaporation rates of the three
stands. The evaporation rate decreases.from the Marsh t0 the
Swamp Shrubs and to the Swamp Conifers. Although the forest
vegetation may be the cause rather than effect of the lower
evaporation, but the higher evaporation rate in the Marsh
may still play an important role in retarding the establish-

ment of forest cover on the area.

109

\

 

 

Plate 1. Set up of the Livingston atmometer in a.
Marsh stand. Inflorescences of Scirpus cypgrinus
can be seen on the upper part of the photOgraph.
Bulb was approximately 18 inches above the ground.

 

 

 

Plate 2. Set up of the Livinyton atmometer in a
Swamp Shrubs stand. Bulb was approximately
18 inches above the ground.

 

111

Weekly averages of temperature and precipitation are
plotted against the average evaporation rate of the three
stands during the 11 week period from July 20 to October 4,
1952, in Fig. 9.

From the graph, we may observe that the evaporation
rate serves as an indicator of temperature and precipitation,
1.9., the higher the evaporation rate, the higher the tem-
perature and the lower the precipitation will be; whereas the

reverse will be true with the lower evaporation rates.

’ 5m

Atlometer Loss... c.c.'s

112

 

(8
s

P

c:

1
Temperature °F.

20.4

 

70 -

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1

l /

v1
0
1

 

   

 

Temperature
----- Evaporation*
--- Precipitation

/' '\

 

.\/ \p//’ l\\ ‘\J/ I j\L//’.’T’.-/"\.\\+O

 

Fig.

20-26 27-2 3-9 10-16 17-23 2h-30 31-6 7—13 1h-20 21-27 28-40

Jul!

29.

Aug. Sept.

Weekly evaporation (Average of three stands) in relation
to temperature and precipitation for the period from
July 20 to October 4. 1952.

* Denotes evaporation of the average of 3 stands.

Oct.

Precipitation-... Inches

113

B.Edaphic factors

A soil is the product of its environment. The most
important-factors in determining the character of the soil
are the climate, living organisms, relief, time and parent
material (lutz and Chandler, 1947). The interrelationships
between the vegetation, soil, and climate are inseparable.
Although climate is most important inkdetermining the range
of a species, or broadly speaking, a formation; the condi-
tion of the soil is often responsible for limiting its
occurrence. In other words, climate determines the climatic
climax of a region. And soil condition within the climatic
region will limit the vegetation either to be that of the
true climax which occurs on the mesophytic, well-drained,
upland soil; or the subclimax (physiographic climax as call-
ed by others) which occurs on either excessively wet or
excessively dry soils. Therefore in the study of vegetation,
soil is one of the most important factors to be taken into
consideration.

The soils in the Higgins Lake area falls into the
Podzol group (Soils and Man, USDA Yearbook 1938). Soils of
three series were encountered in this study. Due to the
excessive moisture contents and the high percentages of
organic matter presented in the soils, only limited labora-
tory experiments were being carried out in the study of

their physical and chemical prcperties.

114

A spade and an auger were used in the field to
examine the soil profiles by way of digging. Soils of all
the quadrats were investigated. As a result, seven soil
samples belonging to five different soil types were en-

countered. They have been coded numerically as follows:

 

 

 

Code No. Soil Type Cover T123
1 Newton loamy sand ggiig-Calamggrostig stand
2 Greenwood peat Scirpus- zyphg stand
3 Rifle peat figlix-Cornug-Algug stand
4 Newton loamy sand ébiggegggi; stand.
5 Rifle peat ghulg-épieg stand
6 Bergland loam Fraxinus-Ulmus-Iilig stand
7 Bergland clay loam 'ngugegggg stand

Soils of same type, such as l and 4, 3 and 5, taken
as samples are due to the fact that different types of
”Vegetation have been recorded.

Samples from depth of six inch intervals were taken
IFor each soil. This demarcation has been set up arbitrarily
because of the lack of zonation in the organic soils. In
Order to obtain a favorable comparison, mineral soils were
Else being divided in the same manner disregarding their

natural depth of horizons .

115
1.5011 profile description

a.Newton loamy sand

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

De th pH Soil Color Soil Texture
Very
4.96 dark Sandy loam
“ brown
6
Dark
4.60 grayish Loamy sand
brown
12“
Grayish or
3.70 dingy white sand
wet_sand
18"
- . -..- -..- 24" _. (Water-table)
Cover Type ----- Marsh (Salix-Calamagrostis
stand)
Physiography --- Flat
t).Greenwood peat
Depth 93 Soil Color Soil Texture
W
4.55 Dark Fibrous
brown peat
6"_.__. (Water-table)
4.43 Brown Fibrous
peat
12"
4.27 Brown Fibrous
peat
18" .
...//"’f-J Cover Type --- Marsh (Scirpus-Iypha stand)

 

Physiography --- Flat

c.Rif1e peat

Depth
I

 

6n

 

12"

 

 

 

18“

 

116

 

 

EH Soil Color Soil Texture
Very
6.21 dark Woody
brown peat
6.09 Dark Woody
brown peat

(Water-table)

5.78 Dark Woody
brown peat

Cover Type ----- Swamp Shrubs

Physiography --- Flat

d.Newton loamy sand

Depth

 

f

 

 

 

 

 

6"
8'0

12“

18‘

 

 

n3 3011 Color 3011 Texture
Very

5.60 dark Woody
brown peat

(Water-table)

6 32 Grayish Wet sand
' brown
Grayish Wet sand
6.35 brown
Cover Type ----- Swamp Conifer forest

of Balsam Fir-Tamarack.

Physiography -—- Flat

e.Rifle peat

 

 

 

 

 

 

Depth pH Soil Color
Very
6.15 dark
brown
60:
Bank
6.08 brown
12“ (Water-table)
Dark
5-32 brown
18"

 

 

Physiography --- Flat

fuBergland loam

 

 

 

 

 

 

 

117

Soil Texture

Woody peat

Woody peat

Woody peat

Cover Type ----- Swamp Conifer forest of
Balsam Fir-N. White Cedar.

 

Depth EH 5011 Color 3011 Texture
1 Very
5.32 dark Loam
" brown
6
5-54 Gray Clay
....... 10" (Water-table)
12'
Gray with Loamy sand
6.00 yellowish with clay
streaks pockets &
8" gravels
l'
’ Cover Tyne ----- Swamp Hardwoods of
' Ash-Elm-Basswood

 

Physiography --- Flat

 

 

 

 

 

 

 

118

 

 

g.Bergland clay loam
Depth EH Soil Color Soil Texture
5.73 Very
° dark Loam
brown
(I!
6.30 Gray Clay
-------- 10'? —— (Water-Table)
12"
6.50 Gray with
yellowish Clay
stains
18“
Cover Type ----- Swamp Hardwoods of
] Elm-Soft Maple

Physiography --- Flat

119

2.8011 volume weight

Lutz and Chandler (1947) have defined 'volume
weight as "the ratio between the dry weight of a given
volume of undisturbed soil and the weight of an equal
volume of water." It is evident that the volume weight is
greatly influenced by soil structure, therefore the deter-
mination should be based on the volume of soil in its
natural field condition.

Soil samples of this study were collected in situ
by means of the.soil core sampler (Cylinder of 3' x 3" in
dimension which gives a volume of 347 cm.). Soil cores of
each horizon were collected in duplicate and placed in ice—
cream containers in order to retain their natural condition.

In the 1aboratory,'a”diSk'ofifilter~paper andzcheeseé
cloth'were.fastenedct010nefend of the"cylinder with a:rubber
band.«They~wbre;weighedvand-oven-dried.at;105°“C.u£or,24—.:
hours and calculation was made on oven-dry basis.

.Results of laboratory determination are shown in
Table 15. Statistical analysis has been made which shows
no significant difference between either different soils or
different horizons of the soil (Table 16).

~ However, from the data, conclusions can be drawn:
(l)The volume weight increases with increasing depth below
the surface; and (2)1ower volume weight in soils of higher

content of organic matter (also see Table 15). These ar‘l

12G

essentially coincideswith-the statement made by Lutz and
Chandler (P. 238, 1947).

Table 15. Volume weight of each of the three
horizons of six soils.

 

 

3011 Type —Code Oven-dry Wt. Volume
No. Horizon in Grams Weight

Newton sand 1 0"-6" 56.5 0.163
6"-12" 143,3 0.413

12“—18" 549.4 1.583

Greenwood peat 2 C"-6" 63.4 0.183
6"-12' 100.3 0.289

12"-18" 105.8 0.305

Rifle peat 3 0"-6" 75.4' 0.217
6..-].2. 114 o} Oo329

12'-18" 129.6 0.373

Newton sand 4 0"-6" 73.8 0.218
6"-12" 568.5 1.638

12"—18“ 601.7 1.734

Rifle peat 5 0"-6" 55.8 0.160‘
6'-12' 61.7 0.177

12"-18" 65.1 0.187

Bergland loam 6 0"-6" (55.6 0.160
, 6"-12" 78.1 0.225
12..“18" 174 oz OoSO}

 

Table 16. Result of the statistical analysis of
the soil volume weight data.

 

ource o egree o um of Mean

_!ariation Freedom Squares Sguare 5_F
Total 17 49893.11 2934.89

Between '

Soil Types 5 23183.78 4636.76 2.89
Between Soil

Horizons 2 10720.44 5360.22 3.35
Error 10 15988.89 1598.89

 

-—o.—

 

121

3.Soil porosity

Most soil porosity measurements are based upon
determinations of volume weight of the soil at some arbi-
trary moisture content (Baver, 1948). In this study, the
total porosity is based on weight of soil at oven—dry, 105°C.,
whereas the separating moisture content between the capillary
and non-capillary porosity is at the tension equivalent to
60 cm. of water (pF 1.6).

a.Capillary, non-capillary, and toatl porosity

Eighteen samples in duplicate were collected in situ
using core sampler. They were taken from each of the three
horizons of six soils.

A disk of filter paper and cheesecloth were fastened
to one end of the cylinder with a rubber band. Cylinders were
then placed into a pan of water such that the water is level
with the top of the soil and allowed to stand for 24 hours.
After the cores were saturated, they were removed from the
pan, let drain freely for one minute, weighed, placed on a
tension table, allowed to drain for 24 hours, and weighed.
Tension tables of four different tensions which have the
equivalent to 10 cm., 20 cm., 40 cm., and 60 cm. of water
were used. Data obtained from the tensions of 10 cm., 20 cm.,
and 40 cm. were used to calculate the percentages of pore
space distribution. Cores were placed on the 10 cm. tension

table first for 24 hours and weighed. The same procedure was

122

repeated for the remaining three tension tables. Cores were

oven-dried after removed from the 60 cm. tension table.
Percentages of the non-capillary, capillary, and

total porosity of the six soils are shown in Table 17.

Table 17. Percentages of soil porosity of each of
the three horizons of six soils.

 

 

 

Code Percentage of Porosity

Soil Typg No. Horizon Non-capillary Capillary Total
Newton.sand l 0"-6“ 37.70 53.66 91.36
6'-12" 13.06 66.94 80.00

l2“-18" 13.78 31.55 45.33

Greenwood peat 2 0"-6" 36.29 56.48 92.77
6“-12" 24.76 63.21 87.98

12"-18" 17.67 67.11 84.78

Rifle peat 3 0“-6" 28.88 53.33 82.21
6"-l2' 15.39 67.46 82.85

12'-l8" 13.04 68.57 81.61

Newton sand 4 o“-6" 39.04 53.03 92.07
6F-12" 16.69 21.26 37.95

12*-18" 8.67 23.49 32.16

Rifle peat 5 08-6” 31.57 59.65 91.22
6'-12" 25.79 62.77 88.56

l2“-l8" 16.92 72.01 88.93

Bergland loam 6 0"-6" 22.74 67.69 90.43
6'-12" 19.42 74.87 94.29

__, 12”=18" 14.58 60.95_ 75.53

 

The data shows that the percentages of non-capillary

porosity are lower than the capillary porosity in all the

soils. This proves the fact that all the soils are poorly-

drained, as Lutz and Chandler (1947) have stated that soils

which possess a low non-capillary pore volume and high capi-

llary pore volume will have high field capacity but the

a.
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infiltration of water will be slow.

123

The high percentages

of the total porosity of all the soils are no doubt due to

the high content of unincorporated organic matter which is

very highly porous in nature.

Table 18. Analysis of variances for non-capillary,
capillary, and total porosity of six soils.

 

 

 

 

 

ggurce of Degree 0? Sum off; Mean 2'
Variation Freedgp_fi Sguares Sguare F ‘_
Non-capillary porosity
Total 17 1483.78 87.28
Between
Soil Types 5 137.78 27.56 1.14
Between Soil
Horizons 2 1104.78 552.39 22.90**
Error 10 241.22 24.12
Capillary porosity
Total 17 4329.78 254.69
Between
Soil Types 5 2671.11 534.22 3.39*
Between Soil
Horizons 2 87.11 43.55 0.27
Error 10 1571.56 157.16
Total porosity
Total . 17 5773 .78 339 .63
Ebtween
Soil Types 5 2221.11 444.22 2.07
Ebtween Soil
Horizons 2 1408.78 704.39 3.28
..Ek-ror 10 2143.89 214.39
*“Denotes significant at 1% level

“Denotes significant at 5% level

124

Results of statistical analysis are shown in Table 18.
There are no siginificant differences between the total
porosity of the six soils. For the capillary porosity,
slight differences (significant at 5% level) exist between
the six soils. Result of the t—Test proves that soils of
code number 2,3,5,6 have higher percentages of capillary
porosity than Newton sand type. The Newton soil has a sub-
soil of wet sand which will certainly be less porous than
the other peat 0r clayey soils.

As for the non-capillary porosity, there are no
significant differences between the six soils. However,
highly significant differences exist between the three
horizons. Results of the t-Test show that the 0"-6" hori-
zons have higher percentages of non-capillary porosity than
the lower two horizons, whereas there is no difference be-
tween the 6"-12" and 12"-18" horizons. This is rather
typical for most of the soils whose sub-soils are less
aerated due to compactness. As in this study, the conti-
1nuous wetness should account for the poor aeration which in
turn explains the incomplete decomposition of the organic

:remains and causes the formation ofpeat

125
b.P0re space distribution

Percentages of pore space for the six soils by
horizons were determined from the volume weight of the soil
at moisture content of the saturated soil core drained for
24 hours under tension equivalent to 10 cm., 20 cm., 40 cm.,
60 cm. of water, and oven-dry conditions. The percentages
of pore space distribution serve to give a better picture
of the pore space under different moisture content.

Results of laboratory determinations are shown in
Table 19.

Table 19. Pore space distribution in percentages
of the six soils by horizon.

   
   
 

--.. ,.___5_ - —- --.r— ...v -»~- -——._. --. ._ --.--. _,____ _. .. _.__..._ -..... ......

 

Tension EQuivalent to cm. of Water
Soil Type_Code No. Horizon 10 cm. 20 cm. 40 cm; 60 cm. Oven-dry

Newton sand 1 0"-6” 21.17 25.22 33.57 37.70 91.36
6"-12" 8.69 9.30 11.38 13.06. 80.00

12"-18" 3.60 4.73 9.54 13.78 45.33

Greenwood 2 0”-6” 21.53 23.66 30.86 36.29 , 92.77
peat 6"-12" 15.30 16.71 21.01 24.76 87.98

‘ 12"—18" 11.18 12.48 15.65 17.67 84.78
Rifle peat 3 o"-6" 22.74 24.09 27.00 28.88 82.21
6'-12" 10.69 11.61 13.66 15.39 82.85

12'-18" 9.10 9.91 11.73 13.04 81.61

Newton sand 4 O"-6" 28.02 31.55 36.73 39.04 92.07
6"-12" 4.21 6.28 13.11 16.69 37.95

12“-18” 3.17 3.40 6.59 8.76 32.16

Rifle peat 5 0“-6" 24.45 26.51 29.93 31.57 91.22
6"-12" 19.31 21.32 24.29 25.79 88.56

12“-18" 11.58 12.97 15.30 16.92 88.93

‘Bergland 6 o“-6' 15.01 16.51 19.45 22.74 90.43
loam 6"-12' 11.30 12.94 16.63 19.42 94.29
12"-18" 10.29 11.21 13.11 14.58 75.53

‘g

 

126

Statistical analysis, as shown in Table 20, indicates

highly significant differences between the different soil

types and between the different soil horizons. The following

statements were drawn from results of the t-Test:

(l)The Greenwood peat and the Rifle peat (Code No. 5)

have higher pore space percentages than the other four soils.

(2)The pore space decreases as depth of the soil

increases.

Table 20. Analysis of variance from data of the pore
space distribution of the six soils by horizons.

 

 

Sources of Degree of Sum of Mean

Variation freedom Squares Sguare_ F
Total 71 5685.87 80.08
Between

Soil Types 5 429.45 85.89 21.00**
Between Diff.

Tensions 3 740.37 246.79 60.34**
Error (a) 15 61.38 4.09 0.18
Between Soil

Horizons 2 3514.58 1757.29 77.79**
Interaction

Between T a H 6 36.42 6.07 0.27
Epror (b) 40 4903.67 22.59

 

**Denoted significant at 1% level.

.>’.v..

 

 

127

4.3011 moisture

In many regions, the occurrence of forest types is
controlled by the supply of water. Generally speaking,
site quality improves with increasing amount of available
soil moisture. However, there is a limit to the amount of
soil moisture which is desirable. Lutz and Chandler (1947)
pointed out if the soil water exceeds a certain limit,
unfavorable conditions for plant growth result because of
deficient aeration and related phenomena.

In this study, the swampy soils are wet throughout
the year or at least for part or all of the growing season.
This makes the accurate determination of soil moisture ip
pipp difficult to accomplish, because most of the methods
in use are not designed to measure soil moisture in very
wet conditions.

Nevertheless, the "electrical resistance method"
(Bouyoucos and Mick, 1940, 1948) was used to determine the
moisture content of two soils. Both of them are Rifle peat,
but the stands are of two different types: a Swamp Conifers,
and a Swamp Shrubs of gpllg-Cornps-Alppg Association. Soils
of these two stands are comparatively drier than soils of
the Marsh and Swamp Hardwoods which at the time of setting

‘up the experiment were both under water-logged conditions.

I)

128
Nylon blocks were buried at 6 inches and 10 inches
depth into the soil column. The lower depth of 10 inches
was selected because the water-table in both the soils were
at about 12 inches. Weekly measurements were taken by means
of the Bouyoucos' Bridge during the period from July 26 to
October 4, 1952. Soil temperatures were also measured at
the same time in order to correct the resistance readings
of the nylon blocks against the room temperature of 76°F.
(Bouyoucos and Mick, 1948).

The soil moisture curves for the two soils, which
have been caliberated in laboratory at room temperature of
76°F. are shown in Fig. 10. (Bouyoucos and Mick, 1940, 1948)

Table 21. Soil moisture data of Rifle peat soils in

two different stand types, obtained during the
period from July 26 to October 4, 1952.

         

 

 

 

 

 

 

 

 

 

éggik —62;Tnche§0 flgoéiich:ig- _%2linche§;ié 10N18c§eiflg
Aug. 2 1.9x103 232 0.92x103 252 2.1x103 199 1.6x103 205
9 4.6x103 209 2.1x103 229 2.9x103 191 2.0x103 200
16 2.9x103 220 1.4x103 240 1.7x103 204 1.4x10: 208
23 2.0x103 250 0.96x103 251 1.5x103 207 1.3x103 210
30 4.2x103 211 2.71103 222 5.41:103 178 4.1xlO3 183
Sept. 6 2.7x103 222 1.2x103 244 2.8x103 192 2.1x103 199
13 5.0x103 207 1.8x103 233 3.8x103 185 3.0x103 190
20 4.0x103 213 2.4x103 225 7.0x103 172 5.03103 179
27 2.2x103 228 0.92x103 252 4.3::103 182 4.011103 184
th. 4 2.8x193 221 1.05x103 24s 5,0x103 179 4.5::103 181
x ----- Corrected resistance reading in ohms.

MC ----- Moisture content in percent.

i"-

129

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130
Weekly data of the moisture contents of the two

soils are shown in Table 21. Graphical presentation is
shown in Fig. 11.

Analysis of variance (Table 22) shows highly
significant difference between the moisture contents of
the two soils as well as between the two different horizons.

Table 22. Analysis of variance of the moisture
contents of the two Rifle peat soils.

 

 

 

Source of ' Degree of Sum of Mean

Variation Freedom uares S uare F
Total 39 21099.78 541.02
Between Dates 9 3143.03 349.22 2.74
Between

Soil Types 1 14478.03 14478.03 113.88**
Error (a) 9 1144.22 127.13
Between Soil

Horizons 1 1600.23 1600.23 193.26**
Interaction '

Between T d H 1 585.22 585.22 70.67**
lgyror (b) 18 149.05 8.28

 

it“Denotes significant at 1% level.

Further t-Test shows that the moisture content of
tdao soils of the Swamp Shrub type (Code No. 3) is signifi-
c=antly higher (at 1% level) than the soils obtained from
tdse Swamp Conifer forest stand (Code No. 5). Whereas in
Ouch soil the lower (10 inches) layer has a higher moisture

'than the upper (6 inches) layer.

 

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Soil Moisture Content in Percent

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240 "‘

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* August

131

 

 

 

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6“ layer of Swamp Conifer stand

——--—12" layer of Swamp Conifer stand
—---'—-—‘7 6" layer of Swamp Shrubs stand
------ “12" layer of Swamp Shrubs stand

Fig. 11. Weekly data of soil moisture content in percent

of the two Rifle pest soils by horizons.

 

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132
5.8011 temperature

Soil temperature is important because it affects
the biological, chemical and physical processes in soils.
Weaver and Clements (1939) have stated that color, texture,
structure, water content, amount of humus, slope and aspect,
and the presence or absence of vegetation cover are among
the factors that directly affect soil temperature. According
to them, water content is the most important because it has
a specific heat about five times greater than the solid
constituent of the soil. Therefore, the poorly-drained
soils, under dense vegetation cover, of flat and low ground,
are colder than sandy or loamy soils of the uplands.

Table 23. Soil temperatures of the two Rifle peat

soils which measured once every week during the
period from July 26 to October 4, 1952.

 

 

 

5011‘““xeeeésature‘ “§L

Date a. Code No. 3; . Code No, 5
July 26 -- 66
Aug. 2 61 61
9 6O 61
16 60.5 60
23 57.5 55
30 60 62
Sept. 6 58 58
13 61.5 62
20 52 53
27 49.5 49

got. 4- 47‘ 46

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133
In this study, soil temperatures have been measured
in only two of the sample plots (shown in Table 23), as a
supplementary procedure in measuring the moisture contents
of the soils. No significant difference has been found
between the two sampling areas, the Swamp Shrub stand and
the Swamp Conifer forest.
6.0rganic matter
The effect of organic matter or humus on forest.
vegetation has been discussed by Toumey and Koretian (1958).
They stated that the humus is a source of supply of both ash
and non—ash nutrients. They also stated that the effect of
organic matter will improve the physical properties of soil.
For example, the incorporated humus improves the soil
structure, increases water holding capacity, decreases the
volume weight, increases the activities of soil organisms,
and has a high exchange capacity; and the unincorporated
organic matter will protect the soil from the compacting
effect of heavy precipitation.
. Decomposition of plant remains is much retarded in
'tho poorly-drained soils duo to the cool climate, excessive
loisture, poor aeration, and unfavorable microbiological
conditions. The litter of the swamp vegetation such as
£1325, ngig, Vaccinium, Lgdgm, 9232;, etc., is highly acid
in reaction which upon accumulation is readily formed into

peat.

134

Braun-Blanquet (1932) has segregated the humus
substances into two qualitatively different groups: the
neutral, mild, saturated; and the adsorptively unsaturated
humus. He stated that ” ----- the interior surface of the
unsaturated humus (of this, peat belongs to) is very great,
involving a large water fixation; hence very acid humus
soils are physically dry."

Twenty-one samples in duplicate from each of the
three horizons of seven soils were used to determine their
organic matter content by the Dry Combustion method.

To 2 grams of finely ground air-dry soil (0.5 or
1 gram for some of the soil whose organic matter is very
high) were added 0.25 gram manganese dioxide and 5 grams
60 mesh carbon free alundum. They were mixed well and
placed in alundum combustion boat and the boat was in-
serted directly into the hot silica tube offurnace, pre-
viously heated to operating temperature of about 950°C.

The flow of oxygen was adjusted to about 100-200 c.c. per

minute. Carbon dioxide evolved in combustion passed thru

'tho purifying section of the train, and was absorbed in a

jpreviously weighed tube filled with ascarite. Grams of carbon

(dioxide evlved was calculated and then converted to percent

of organic matter by using the converting factor 0.471.
Results from the laboratory determination are shown

in Table 24.

135

Table 24. Organic matter in percent of seven soils
by horizon as determined from Dry Combustion

 

 

 

method.

Organic Matter in Percent

Code ; . Replications
Soil Type__ No. Horizon I _, II Average
Newton sand 1 0'-6” 59.24 40.38 49.81
69-12" 16.49 18.90 17.69
12“-18" 1.52 1.00 1.26
Greenwood peat 2 0i-6" 67.68 66.62 67.15
6"-12" 78.08 71.98 75.03
12'-18" 33.77 50.29 42.03
Rifle peat 3 0'-6"‘ 51.91 48.75 50.33
6"-12“ 63.23 53.32 58.27
12"-18" 41.40 44.373 42.88
Newton sand 4 0'-6' 47.82 48.55 48.19
6“-12" 0.79 1.30 1.04
12'-18" 0.99 0.79 .0.89
Rifle peat 5 0“-6' 42.41 46.01 44.22
12"-18" 40.40 43.38 41.89
Bergland loam 6 0“-6" 46.12 30.47 38.30
6"—12" 32.24 21.57 26.91
12”-18" 1.33 1.82 1.58
Bergland 7 "-6“ 38.49 42.14 40.31
clay loam 6"-12' 3.72 3.37 3.55
_p 12"-18" 1.35 1.29 1.32

 

 

 

 

Statistical analysis which is~ shown in Table 25

shows that there are highly significant differences (at 1%

‘level) between the organic matter of the different soil

‘types, different soil horizons, and also the interaction

of soil type and horizon.

136

Table 25. Analysis of variance from the data of

organic matter of seven soils by horizons.

 

 

Source of Degree of Sum of Mean

Variation Freedom Squares. Sguare F
Total 41 22913.62 558.87 105.85
Between

Soil Types 6 12168.12 2028.02 75.42**
Between

Replications 1 27.52 27.52 1.02
Error (a) 6 161.31 26.89
Between Soil

Horizons 2 6135.05 3067.52 105.85**
Interaction

Between T d H 12 4015.95 334.66 11.55““
Error (9) 14 405.67 28.98

 

*“Denotes significant at 1% level.

The following observations are drawn from tho t-Test:

a.For different soil types h-—-- The Greenwood peat has

the highest organic matter content, followed by the two Rifle

peat types. Next comes the two Newton sand types and both the

Bergland soils.

The high percentages of organic matter in all the

{peaty soils can well be expected as the result of vegetative

accumulation in the peat. The greater amount of coarse woody

materials in the Rifle peat should account for the lower

organic matter content than found in Greenwood peat which

is composed of mostly finer textured materials.

The lower organic matter content of the Newton and

IBergland soils is due to the fact that they are mineral

137

soils whose substrata are low in organic matter, in spite of
the high content of organic matter in the t0p layer of the
two soils.
b.F0r types at same horizon

(l)O"-6" Horizon ----- Greenwood peat has the highest
organic matter; and the Bergland loan has the lowest. The
rest of the soils are in between without differences.

(2)6"-l2" Horizon ----- Organic matter in soil code
No. 2 3 5. 6,1 4,7. The higher organic content in the
first three soils is attributed to the characteristic of peat
which is naturally high in organic matter. The decreases in
organic content of these soils account for the fact of vege—
tational changes from Marsh to Swamp Shrubs and to Swamp For-
est vegetations. The lower organic matter content of the
Newton and Bergland soils is due to the mineral origin of
the subesoil. The differences between them are probably due
to the variation in thickness of organic surface soils.

(3)12'-18' Horizon ----- Results of the t-Test show
that the three peat soils are higher in organic content than
the Newton and Bergland soils. This coincides with the dif-

Irerence between the pest and mineral soils.

138

7.Soil reaction (pH)

Most forest soils have an acid reaction. Acidity of
the soil may affect tree growth. However, certain species
particularly of the family Ericaceae, growbest on acid soil.

Under conditions of poor drainage, highly acid con-
ditions are apt to develop. Weaver and Clements (1938) stated
that ”In soils of organic origin, basic salts are present
in only small amount. The soil develops more or less marked
acidity as the result of the accumulation of humus under
conditions of poor aeration and sometimes by the setting
free of acid from the mineral constituents of the soils."

The Beckman pH meter was used in the determination
of the soil acidity in this study. Twenty-one samples in
duplicate were tested. For the organic soils, soil-water
ratio of 1:10 by weight, and 5 grams air-dry soil samples
were used. For the mineral soils, 10 grams of soil at soil-
water suspensions of 1:5 by weight were employed for the
determination. .The higher soil-water ratio for the organic
soils was necessary to make better contact between the soil
iamd the glass electrode. The use of a small amount of
<3rganic soil for samples was made possible by their high
13ercentages of organic content.

A series of soil samples were weighed up.and to such
sample the required amount‘ofrwster weseddedw Thor soil sus—
'Dénsioasawore stirred for'one.minuteg and the pH determined.

Laboratorygresults are shown in Table 26.

159

Table 25. Soil reaction (pH values) of the seven
soils by horizon.

 

 

 

 

 

 

 

Code - 5gplicationg

Soil Type_ No. Horizon I II ' Average
Newton sand 1 0"-6" 4.98 4.95 4.96
6"-12“ 4.65 4.55 4.60
12"-18' 3.75 3.65 3.70
Greenwood 2 0"-6" 4.55 - 4.55 4.55
peat 6"-l2" 4.48 4.38 4.43
12"-l8” 4.22 4.32 4.27
Rifle peat 3 0"-6” 6.20 6.22 6.21
6"-12" 6.06 6.12 6.09
12"-18' 5.78 5.78 5.78
Newton sand 4 o“-6“‘ 5.65 5.56 5.60
6“-12" 6.40 6.25 6.32
12"-18" 6.45 6.25 6.35
Rifle peat 5 0"-6" 6.15 6.15 6.15
6"-12“ 6.02 6.15 6.08
12"-18" 5.35 5.28 5.32
Bergland loam 6 0“-6' 5.75 5.70 5.73
6"-12" 6.25 6.35 6.30
12"-18" 6.40 6.60 6.50
Bergland 7 0"-6“ 5.25 5.20 5.23
clay loam 6”-12" 5.52 5.55 5.54
. 12"-18" 5.85 6.15 6.00

 

Statistical analysis has been carried out for five

of the seven soils tested. The arbitrary horizons of 6 inches

interval in this study are set up primarily for the organic

soils (Azonal) which are undifferentiated in natural horizons.

In this experiment, if both the Bergland soils are

thrown in for statistical analysis, erronous results would

9‘"

140
be produced because of the different thickness in the
correspoding horizons of these two structurally similar
soils. Therefore, only one of the two is being analyzed.

The same applies to the Newton soils.

Table 27. Analysis of variance for the soil
reaction data of five soils by horizon.

 

 

 

‘ource of Degree of Sum of Mean

Variation Freedom Sguares Sguare F
Total 29 2200.97 75.90

Between , .

Replications 1 0.04 0.04 0.11
Between

Soil Types 4 1848.14 462.04 l248.76**
Error (a) 4 1.46 0.37

Between Soil '

Horizons 2 112.27 56.14 93.57**
Interaction

Between T e H 8 233.06 28.51 47.52**
Error (b) 10 6.00 0.60

 

‘senenotes significant at 1% level.

Analysis of variance in Table 27 shows a highly
signifiCant difference between the five soil types as well
as between horizons. The t-Test gives further results as
follows:

a.Eor different soil types ----- The two soils of the
Open-meadow type (Greenwood peat and Newton sand) were
strongly acid in reaction, followed by the two Rifle peat
soils. No difference was found between the former two soils,

whereas the Rifle peat of the Swamp Conifer Forest stand was

 

 

141

slightly more acid than the soils of the Swamp Shrub type
(significant at 5% level). The Bergland soil of the Swamp
Hardwood stand was the least acid and almost neutral in
reaction.

b.For different horizons ----- The l2“~18" horizon
was significantly more acid_(at 1% level) than the top two
layers, while there was no difference between the latter
two horizons.

This was true for the organic soils and was probably
due to the excessive moisture present in the lower layer
which in turn caused poorer aeration and less decomposition
of plant remains.

As for the Bergland soil, the pH value increased
with increasing depth. This is typical to podzel soils as
stated by lutz and Chandler (1947): .

“The H layer or the A horizon in odzols is
commonly most acid and the maximum values are
commonly encountered in the lower part of the
B horizon."

A- composite summerization of all the edaphic

characteristics for the seven soils by horizon is shown

in Table 28.

Table 28. Composite summation of edaphic

Soil Type

Newton sand

Greenwood
peat

Rifle peat'

Newton sand

Rifle peat

Bergland
loan

Bergland
clay loam

characteristics of seven soils by horizon.

Code

Capil—

142

Porosity in Percent Organic

Volume Non-Ca- Matter.

pH

No. Horgzon Height pillar: lary Total percent

 

1

0"-6'
6"-12“
l2"-l8”

077-6'.‘
6“—l2“
12'-18'

08-6"
6"-l2'
l2"-l8"

0.8-6'.
6“-12'
l2"-18'

Ou_6u
6"-12"

'12'-18"

0.163
0.413
1.583

0.183
0.289
0.305

0.217
0.329
0.373

0.218
1 o638
1.734

0.160

0.177
0.187

37.70
13.06

13.78

36.29
24.76
17.57

28.88

15.39
13.04

39.04
16.69
8.67

31.57
25.79
16.92

"‘ -----

53.66
66 o94
31-55

56.48
63.21
67.11

53.33
67.46

68.57

53.03
21.26
23.49

59.65
62.77
72.01

91.36
80.00

45.33

92.77
87.98
84.78

82.21
82.85
81.61

92.07
37.95
32.16

91.22
88.56
88 o9}

49.81

17.69
1.26

67.15
75.03
42.03

50.33
58.27
42.88

48.19
1.04
0.89

44.22
44.15
41.89

38.30
26.91
1.58

4.96
4.60
3.70

4.55
4.45

4.27

6.21
6.09
5.78

5.60
6.32
6.35

6.15
6.08

5.32

5-73
6.30
6.50

5.23
5.54
6.00

143

C.Biotic factors

In this paper, biotic factors are considered under
three groups: (1)1nsects and diseases, (2)Animals, and
(3)Men. No attempts have been made toward detailed studies
of these factors because they were not believed to signifi-
cantly influence forest succession on the poorly-drained
soils of this region. Onlthhose observed to be relatively
important were covered.

l.Insects and diseases

Generally speaking, most of the insects and diseases
have species preferences or at least do not injure all species
equally. This may be due to the composition and age-pattern
of the forest. Therefore swamp forests which are mixed in
composition and uneven-aged in most cases probably will not
be subjected to great insect damage.

Some of the insects and diseases have had their
epidemic periods. However, unusual damages were not reported
in this region. Some of the common ones have been observed
‘but were not to be considered destructive, at least from the
ecological standpoint.' Important ones encountered in the
field were:

a.Insects
(1)8pruce budworm (Choristoneura fumiferana Clem.)
The spruce budworm is native to North America.

Destructions caused by this pest to the spruce and fir have

144
been great according to Brown, MacAloney, and Dowden (USDA
Yearbook, 1949). Outbreaks of this insect has been recorded
in Maine in 1910, in Minnesota in 1913, and more recently
in Canada in 1935 and 1944. No report of unusual abundance
of the budworm has been received from the Lake States.
However, light defoliations caused by this insect has been
noted in the Higgins Lake region.

Upon the study of the biology and feeding habit of
the spruce budworm, white and black spruce appeared to suffer
less from attack than red spruce and balsam fir. And because
spruce is far more valuable for lumber and pulp than balsam
fir, it has been cut more heavily and its proportion in the
forests has thus been reduced. Balsam fir is far more aggres-
sive than spruce in seeding-in after logging operation, fire,
or wind damage. Man's activity and the spruce budworm, there-
fore have often contributed to a gradual conversion from a
forest containing a high percentage of spruce to one in which
balsam fir predominates and which is far more favorable for
the spruce budworm.

Three silvicultural practices can be done to increase
the resistance of the forest to spruce budworm attack: (a)To
clear cut mature and over-mature balsam fir stands, (b)To
Operate balsam fir stand on a short rotation, and (c)T0 try

to increase the proportion of spruce in the stand.

145

(2)Larch sawfly (Iygaeonematus erichsonii)

I The larch sawfly is the most important sawfly to
attack conifers. From time to time, it appears in destructive
numbers, defoliating extensive areas of tamarack, and it may
also attack some other trees and shrubs.

Westveld (1949) and others have stated that the
tamarack type was almost completely destroyed by this insect
many years ago. In this area, some of the tamaracks in the
Swamp Conifer Forests are the victims of this_pest.

According to Doane (1936): "Little can be done toward
controlling this insect in the swamp areas. ----- Importation
of one of the European parasites has been evidently proved
successive in Manitoba (Watson, 1931).“

b.Diseases -

The important diseases pertaining to the swamp forests
can be grouped into decay (or rots) and cankers. Fungi are
the causal agents for both groups. In addition to the loss
of merchantAbale timber, trees weakened by the rots or cankers
are much more easily broken off and therefore these diseases
indirectly disturbs the original stand characteristic and
composition.

The common diseases either being observed or believed
to be present in the swamp forest of this region are listed

by their causal agents and the host Species:

Ogcanism
Armillaria melea

Formes igniarius
Formes pini
Polyporus balsameus

Polyporus schweinitzii
Perla subacida

Stereum sanguinolentum

Hypoxylon pruinatum

Nectria sp.

2.Animals
In general, the
wildlife. When there is
wildlife population and

146

Damage caused by organism and host species

Occasionally cause rot in black spruce,
yellow birch, and American elm.

lost important in aspen, also attacks
yellow birch, to cause white heart rot.

Cause red ring rot in tamarack and
black spruce.

Cause brown butt rot in balsam fir,
northern white cedar, and hemlock.

Cause red brown butt rot in black spruce.
Attacks balsam fir especially.

Cause red heart rot in balsam fir
and spruce.

Common and sometimes fatal in aspen.

Common in yellow birch, occasional in
red maple, basswood, and American elm.

forests provide food and cover for
a reasonable balance between the

food supply from the forests, there

will be no damage to the forests. However, repeated browsing

by over-p0pulated wildlife definitely damages reproduction.

Swift (1949) reported that reproduction used most heavily as

food by deer was that of maple, oak, northern white cedar,

basswood, ash, birch, elm, etc., with prevalence in that

order. Of these, northern white cedar, ash, elm, and maple

are the important components of the swamp forests.

147

3.Men

The effect of man's destruction upon vegetation is
visible everywhere. The most important associates of man
in the destruction of vegetation are "fire" and "cutting".
Although fire occasionally may be caused by lightning by‘
far the greatest number are man caused.

In the early lumbering period, general cutting
practice was clear-cutting. Actual evidence supports the
fact that in certain sections of the state fires destroyed
more merchantable‘timber than was out during that period.
However, the swamps have not been subjected to burning
nearly so often as the uplands. This may attribute to the
fact that the swamps are usually wet with less chance for
fire to occur. This does not mean there are no fires in the
swamps. As we can see during the dry season, the water-table
in a coniferousswamp may drop to a low level and this plus
the fact that the litter of the swamp stands become readily
flammable, may result in fire doing great damage.

Clear-cutting was the common practice in early
lumbering period. This will eventually result in a shrubby
growth on the site before the reproduction of the original
Species can re-establish; and this may not be accomplished

for a considerable long period of time.

148

Unfavorable cutting sometimes may alter the stand
composition, as for 'example, when selective lOgging is
employed to remove valuable large black spruce from a coni—
ferous swamp. This will result in the stand being dominated
by inferior species such as balsam fir and culled northern
white cedar. Eventually the unsuitable cutting operation
will disturb the natural sequence of succession either pro-
gressively or retrogressively.

Secondary succession begins when the original stands
are burned or otherwise denudated through some other agents
such as destructive cutting, epidemic insect or disease
damages etc.. Generally speaking, on lightly burned areas,
original species such as black spruce or balsam fir of the
Swamp Conifers may easily be reestablished in a short time
because of favorable regeneration characteristics. On severely
burned areas succession may initiate from the sedgeegrass
stage, the swamp shrub growth of figlingornus-Alngg associa-
tion, or the Lowland Aspen type. In either case, the sub-
climax type of Swamp Conifers or the Bwamp Hardwoods will
replace the temporary types depending upon the original
composition and the nature of the soils. On shallow peat
with mineral substratum, Swamp Hardwoods will have better
chance in succession; whereas on bog or peat soils, Swamp
Conifers probably will succeed. Repeated fire generally
favors the continuance of Aspen type (Gates, 1930).

FOREST SUCSESSION

Cooper (1913) has divided succession into a series
of two types, the xerarch and hydrarch. Hydrarch succession
initiates from hydrophytic habitats such as lakes and ponds
and becomes more and more mesephytic in its successive stages.
This, the hydrarch succession, is the type to be discussed
in this study.

Five vegetation types were found on the poorly-drained
soils in the Higgins Lake area. They were the Marsh, the Swamp
Shrubs, the Swamp Conifers, the Swamp Hardwoods, and the Iow-
land.Aspen. A diagram showing successiohal relationships be-
tween the different vegetation types is shown in Fig. 12.
Trends of succession are based on the characteristics of the
soils and the occurring vegetation. No attempts were made to

study the succession prior to the Marsh type.

A.The Marsh (or Open-Meadow type)

Marshes were found on two types of soils, the boggy
Greenwood pest and the Swampy Newton loamy sand. These two
soils have the common characteristics of being strongly acid
in reaction; but they do differ in moisture and organic con-
tents.

The Greenwood peat is comparatively wetter than the

Newton loamy sand because the former has a lower volume weight

 

 

150

Hemlock-White Pine-Northern Hardwoods
(Upland Climax Type)

\

\

Swamp Hardwoods of
Fraxinus-Ulmusrloer
(Subclimax 4LType)

Swamp Conifers of

Thuja-Picea-Abies

(Subclimax Type)
/

1"-

\

’./
Lowland

Larix

 

 

 

v Swamp Shrub
Denudation of , /"

Swamp Forests ,,’~
by destructive
cutting, fire, ‘ \
& other causes. '“~\,

 

 

 

 

Fig. 12. Diagram showing successional relationships
exhibited between the associations of poorly
drained soils in the Higgins Lake area.

An arrowhead indicates the direction of succession.

151
and higher capillary porosity, which indicates a higher
water holding capacity. This wetter condition can also be
verified by the fact that the water-table in the Greenwood
peat is very near to the surface as compared to the lower
water-table, at about twenty-four inches, in the Newton
soil during the drier season.

The organic content of the Greenwood peat is con-
siderably higher than that of the Newton sand. This should
be as eXpected due to the difference between organic and
mineral soils. The shallower organic layer on top of the
wet sand of the Newton soil indicates its better-drained
condition than the Greenwood peat which has a deep peat
soil column.

The difference in vegetation components is another
indication that the vegetation on the Newton soil belongs
to a later stage. The plants found on the Greenwood peat

are more hydrOphytic such as Sggittaria, Alisma, Juncus,

 

Eleocharis, Typhg, and the abundance of §igm suave and
Cicuta bulbifera. On the Newton soil there is a more luxi-
rant growth of Sglgg and the presence of such shrubs as
Alggg, S iraea, and nggg,

Continuous filling-in onto the harsh site by the
mat forming plants such as sedges, rushes, grasses, cat-
tails, etc., will result in one of two possibly successional

stages:

(K

 

 

152
l.The Swamp Shrubs, or the Swamp Conifers will succeed
on the poorly-drained boggy sites.
2.The Lowland Aspen may succeed on relatively better
drained sites as a temporary type and later give way to the

Swamp Hardwoods or the Swamp Conifers.

B.The Swamp Shrub type
As the sedge mat gradually invades the poorly-drained
peat, shrubby species will invade the Marsh to form a shrub

stand which is principally composed of Species of Salix,

 

Alpgg, and Cornus.

Plants such as Aronia nigra, Potentilla fructicosa,
Betula pumila, Spirggg alba, and Rubus hispidgg etc., are
the indicators of the bog sites. Consequently, the Swamp
Conifers of ggpea-gbieg, or ngig, or a mixture of these
species will succeed this type in a longer or shorter
period of time.

,However, the Swamp Shrubs may also give way to the
Swamp Hardwoods of Fraxinus-Ulmggngggg community on the

better drained sites, or on soils of mineral origin.

153
C.Swamp Coniferous Forest

The principal tree species of this type are Thpjg
occidentalis, Picea mapgana, Abies balsamea, and ngig
laricina. The Thglg, ggpggJ and L321; may occur in almost
pure stands, on the other hand, they may occur in mixtures
of any prOportions.

This type may directly replace the Marsh vegetation
whenever it is able to become established. However, it is
more apt to succeed the Swamp Shrub type.

_ L531; is a rapid growing, short lived, deciduous
coniferous species. It usually invades the cpen, burned over
boggy sites as a pioneer species due to its shade intolerant
character. Generally the ngig forms a pure stand, and an
abundant growth of shrubs usually is found in the stand be-
cause they are less densely shaded. In succession, this type
will be followed by other tree associations which are able
to shade it. The usual ones are the giggg marigpa on very
wet sites, or the Thglgegigggrépigg_type on comparatively
drier sites.

The black spruce (giggg mariana) is the most common
tree species to follow the Swamp Shrubs or the ngig in
succession due to its ability to withstand shade, excessive
soil moisture, and soil acidity.It usually associates with
balsam fir and northern white cedar and later as the stand
advances will be replaced by the northern white cedar (Gates,

1942).

u.-

154

The Swamp Coniferous forest, especially the Thuja,
when left undisturbed and fully developed, will attain a
very dense growth so that almost no ground vegetation grows
in its shade. However, in openings, a great number of plants
of different genera do occur.

Th2}; is a subclimax typo in the region, considering
the Hemlock-White Pine-Northern Hardwoods being the climax
type on the well-drained, mesophytic, uplands. Without pro-
nounced changes in drainage, Thuj; would probably maintain
its composition for some time to come. As Gates (1942) has
stated the 1334; association is the subclimax type in boggy
areas. He pointed out that the ultimate outcome would be the
upland vegetation if the addition of soil material or a
lowering of water-table converts the site to that of the
upland.

Zassda (1952) in study the reproduction on cut over
swanplands in the upper penisula of Michigan, has come up
with the conclusion that " ----- the important species in the
peat swamp was black spruce, but after logging reproduction
of black spruce is poor. Reproduction on the muck soils con-
tains much less northern white cedar than the original stand.
On the wet mineral soils, the stocking of conifers is decided-
ly lower than the original forest. ----- Hardwood invasion is

common on all swamp soils. ----- '

155

Westveld (1948) stated that the black spruce and
northern white cedar are suffering from competition of
hardwoods when the latter do appear in the composition
because of the fast growth of the hardwoods in the early
stage which more or less supresses the reproduction of the
conifers.

These lead me to the believe that, on the Swamp
Conifer sites, as time goes on, the Swamp Hardwoods, pro-
bably the Fraxinus-glmggyépgg community, will eventually
replace the original conifers before reaching the meso-
phytic climax type of the upland.

Results from the vegetation analysis of the Swamp
Conifers (Table 4) show that the balsam fir is the prede-
minating species, followed by the northern white cedar,
tamarack, black spruce, red maple, and yellow birch in that
order. Phytographs of these dominant species are shown in

Fig. 13.

156

a,

 

 

<7
o

Balsam Fir N. White Cedar Tamarack

 

e
0

Black Spruce Red Maple

®

Yellow Birch Black Cherry

OA --- Percentage of relative density

OB --- Percentage of total frequency

CC --- Percentage of Size Class distribution
OD --- Percentage of relative basal area

Fig. 13 Phytographs of the important species
of the Swamp Coniferous Forest type.

157

D.Swamp Hardwoods of Fraxinus-Ulmus-Acer association

The Swamp Hardwoods in this region occurs commonly
on those swamp areas that are occupied by soils which have
a better decomposed, relatively shallow, peaty organic layer
on a mineral substratum. These soils are better-aerated and
less acid than those of the bog sites. The stands contain
species of both the swamp conifers and the upland hardwoods,
generally with the hardwood species dominating.

Table 29. Data of relative feequency, relative

density, and relative basal area of the dominant
components of the Swamp Hardwoods.**

 

 

 

Si;:i§§==========:"_1: Re1.Freg. Rel.Den. Rel::S.A::=IT7fi
Black and Green Ash 21.0 44.7 27.0 92.7 .
Red and Silver Maple 13.2 11.9 33.3 58.4
American & Slippery Elm 18.4 9.7 28.6 56.7
Northern White Cedar 7.9 16.0 10.3 34.2
Balsam Fir . 7.9 8.2 3.2 19.3
Yellow & Sweet Birch 7.9 2.5 2.8 13.2
Basswood 5.4 3.3 2.7 11.4
Hemlock 2.6 0.5 0.4 3.5

 

*‘Produced from Table 2.

*Designates Importance Value which is the sum of
relative frequency, relative density, and relative
basal area. ‘

Table 29 shows the result of a vegetation analysis

of the Swamp Hardwoods. The ashes (Black and green) are

 

 

158
predominating, followed by the maples (red and silver), the
elms (American and slippery), northern white cedar, balsam
fir, the birches (yellow and sweet), basswood, and hemlock
in that order. Phytographs of these dominant species are
shown in Fig. 14.

From the standpoint of soil development, the Swamp
Hardwood is more apt to be an advanced type that succeeds
the Lowland Aspen or the Swamp Shrubs of the relatively
better-drained mineral soils of the river and stream banks.
In many cases, the Swamp Hardwoods are found on depressions
or flat areas bordering the swamps and bogs and connected
on the other side to the uplands. Therefore, geographically,
it occurs on sites which are intermediate between the swamps
and the uplands. Whereas from the standpoint of the vegeta-
tion itself, the Swamp Hardwood type is composed of both the
swamp coniferous species and the upland hardwood species
which shows a true transition between the Swamp Conifers and
the Hardwood Upland.

Gates (1942) pointed out that " henever boggy or wet
ground has been cut over or burned over there is a possibi-
lity of the development of the lowland forest association of
Fraxigps piggg and Apgg_rubrum. Only those of Thglg.gccid§p-
351;; are expected to result in succession, unless the area
is being changed from a lowland to upland. In the latter case
the upland hardwood species more frequently invade and attain

dominance."

 

ifi

Black Ash

 

159

Red Maple Green Ash

 

N. White Cedar Slippery Elm American Elm

O V

Balsam Fir

OA --- Percentage

OB --— Percentage
OC --- Percentage
OD —-- Percentage

Basswood Yellow Birch

of relative density

of total frequency

of Size Class distribution
of relative basal area

Fig. 14 Phytographs of the important species of the Swamp

Hardwoods type.

160

Braun (1950) has discussed the Swamp Hardwoods as the
river swamp community occurring on the wet but better drain-
ed habitat, and stated that on better-drained sites, a few
additional hardwood species may enter into the present Elm-
Ash-Maple communities, suggesting its replacement under good
drainage conditions by mesophytic hardwood forest.

It is probable that the Swamp Hardwood type will

succeed the Lowland ASpen or the Swamp Shrubs of Salix-Cornus-

 

Alggg type on better-drained mineral soils of the swamps
(Gates, 1942). And it may also invade the Swamp Conifers of
Egglg-Apiggeglggg stand when the drainage improves and soil
reaction is not too acid.

The Swamp Hardwoods type is a subclimax on the poorly
drained sites of the region. Without pronounced changes in
climate and drainage, this type will perpetuate and remain
for a relatively long period of time before the upland hard-

woods can be established.

161
E.The Lowland Aspen type
This type is characterized by the dominance of
Pppulus tremulques, the trembling aspen. Inception of this
type can be of two possibilities:
l.It succeeds the Marsh on the relatively better-drained
soils, as a temporary type. After a relatively short period
of time it will give way to either the Swamp Conifers or the
Swamp Hardwoods.
2.It is established as a temporary fire type after an
area of swamp forests, most commonly the Swamp Conifers, is
burned over. Gates (1942) stated that an ASpen association
developing on an area following one or two fires usually
givesplace to a higher genetic association in a comparatively
short time. While further fires favor the continuance of the
Aspen association. He pointed out the Thglg association may
enter immediately and replace the Aspen association in twelve
to twenty years, or a mixture of spruce-balsam fir will
succeed in an even shorter time. The Swamp Hardwoods of Ash-
Elm community may also replace the aspen sometimes.
Usually the original type previously occupying the

site will replace the aspen in scoondary succession.

SILVICULTURAL CONSIDERATIONS

Five types of vegetation were recognized on the
poorly-drained soils in the area: The Swamp Hardwoods, the
Swamp Conifers, the lowland Aspen, the Swamp Shrubs, and the
Marsh. Silvicultural treatments are here considered only
for the former three types.

Since both the Swamp Hardwoods and the Swamp Conifers
are the physiographic, subclimax type, they are able to per-
petuate easily if they be left undisturbed and without pro-
found changes in environment. However, men are? interested in
growing and harvesting the desirable species for use, suitable

silvicultural treatments must be adapted in order to do so.

A.Swamp Hardwood type

The Swamp Hardwoods type is being recognized as the
'most advanced forest type on the poorly-drained soils in the
region. The major species of the type were the American and
slippery elm, red or silver maple, black and green ash, with
some associated species of basswood, northern white cedar,
yellow birch, among others. Due to the excessive moisture of
the soil, these species were not as well deve10ped as those
on the upland sites. At the present time, most of the stands
were uneven-aged, mature, and well stocked with advance growth

of red maple, ash and elm.

163
In the past, logging was carried out on a selective basis to
remove merchantabale and desirable Species. Due to the fact
that this type occurs mostly in small tracts bordering larger,
conifer swamps, intensive silviculture will probably not pay.

Westveld (1949) suggested clear cutting followed by

improvement cutting or thinning for this type. It was based
on the easiness of the establishment of reproduction. More
recently, Eyre and Zilligitt (1953) proposed partial cuttings
for Northern Hardwoods in the Lake States which I believe can
well be applied to the Swamp Hardwood type. However, due to
the shallow root systems of some of the species, resulting
from excessive soil moisture, only light partial cuttings

should be employed.

B.Swamp Conifers type
The Swamp Coniferous stands occurred both as even-
and uneven-aged in the area. Due to the generally high den-
sity of the even-aged trees, there was a wide variation in
the size of individual trees, giving an uneven—aged appearance.
The principal Species of this type were the black
spruce, balsamfir, and northern white cedar. Black spruce
and balsam fir are chiefly used for pulpwood, and the northern
white cedar is mainly used for poles,posts and shingles.
The associated species of tamarack and hardwoods have little

commercial value..

164

lebarron (1948) suggested clear cutting in strips,
the removal of 50 to 75 feet strip along the leeward side of
the stand at l to 3—year intervals, for the even-aged, thrifty,
black Spruce stands. As for the defective, over—matured stands,
clear cutting was suggested. He stated that reproduction is
usually adequate, and it can be improved by slash disposal as
piling to expose more soil surface, and by prevention of logg-

ing damage to advance reproduction.

C.Lowland Aspen type

This is a temporary, or fire type on the poorly-drained
sites. In this area, the stands were even—aged and mostly
under-stocked.

Eventually either the Swamp Conifers or the Swamp
Hardwoods will replace this type depending on the available
seed trees near by or the advanced reproduction already on
the site.

The Aspen usually lasts for a single generation be-
cause it is very intolerant and easily susceptible to disea-
ses. Therefore, on stands having advanced growth, clear cutt-
ing is probably the only method to regenerate this type.
Whereas on stands lacking desirable advanced reproduction,
conversion of the stand to that of desirable Species by clear

cutting and planting was suggested by Westveld (1949).

SUMMARY

The Higgins lake area is located in the northern cen-
tral part of the lower penisula of Michigan. Ecologically,
it is within the Hemlock-flhite Pine-Northern Hardwoods Forma-
tion as defined by Nichols (1935), Braun (1950), among others.

General description and history of the area is review-
ed. Climate, geology, soils, and cover types of the area are
also discussed.

Five types of vegetation were recorded on the poorly-
drained soils in the area. They were: (l)The Marsh or Open-
meadow type, (2)the Swamp Shrubs of the Sglig—Cornus-Alggg
Association, (3)the lowland Aspen of Pogulus-_§;ig Association,
(4)the Swamp Coniferous forest, and (5)the Swamp Hardwood
forest. Vegetation inventory was carried out in the summer
of 1952. Quadrats of three sizes were used for the vegetation
study, i.e., 1/5 acre quadrat for tree vegetation; l/lO acre
quadrat for shrub vegetation; and l/lOOO or milacre quadrat
for vegetation below 6 feet tall, the herbaceous layer.

Ecological factors were considered in three groups:
(1)Climatic, (2)Edaphic, and (3)Biotic. Among the climatic
factors discussed, temperature, relative humidity, precipita-
tion, and evaporation, evaporation is believed to show the
combined effect of the other three. Data of evaporation rates
in three different vegetation type were obtained by using

Livingston atmometers. Results show that the evaporation

166
rate decreases from Marsh to Swamp Shrubs and to Swamp forest.
This indicates that although the forest cover may be the cause
of the lower evaporation, but the higher evaporation rate in
the Marsh may still be an important factor in retarding the
establishment of forest cover on that area.

Soils of all the quadrats were investigated in the
field, seven soil samples belong to three soil series were
encountered and were taken into laboratory. They belonged to.
five soil types: the Newton sand, Bergland loam, Bergland clay
loam, Rifle peat, and Greenwood peat. On each of the two sam-
ples of the Newton sand and the Rifle peat, different types
of vegetaion were found. Samples of each soil by horizon were
collected to determine their characteristics. The following
properties were determined in laboratory experiments: (1)8011
volume weight, (2)8011 porosity, (3)0rganic matter content of
the soil, and (4)3011 reaction. Soil temperature and soil
moisture were determined in the field. Statistical analyses
were used to analyze all the laboratory data which give the
following results:

l.No differences between the volume weight of the soils.

2.No differences between the total porosity of the soils.

3.The Newton sand had a lower capillary porosity than the
other soils.

4.3011 moisture content is believed to be very high in the

Marsh type which has a water-table near the surface. The

167
available data from the field determinations for the
two vegetation types show that the_moisture content in
the Swamp Shrubs type was higher than in the Swamp Coni-
for type. This leads to the belief. that soil moisture
content decreases from the Marsh to the Swamp Shrubs and
to the Swamp Forest. 1 1
5.5011 temperature was relatively constant between the
Swamp Shrubs type and the Swamp Coniferous forest.
6.The peaty soils had higher organic matter contents than
the mineral soils, with the Greenwood peat of the Marsh-
type having the highest percentage.
7.Soils of the Marsh type were strongly acid in reaction,
followed by the two Rifle peat soils of the Swamp Shrubs
type and the Swamp Coniferous forest. The Bergland soil
of the Swamp Hardwood type was the least acid and almost
neutral in reaction.
Biotic factors are discussed under three groups: (1)
Insects and diseases, (2)Animals, and (3)Men. Biotic factors
are not believed to be of great sifnificance to influence the
forest succession on the poorly-drained soils in this area.
The most important one is probably the fire which will induce
secondary succession.
Diagram of forest succession on the poorly-drained
soils in this area is shown in Fig. 14. The principal trend
of succession is believed to be that from Marsh to Swamp

Shrubs type to Swamp Conifers to Swamp Hardwoods and finally

168

after a relative long period of time to the upland, meso-
phytic, climax type of the Hemlock-White Pine-Northern Hard-
woods. The lowland Aspen type is believed to be the

fire or temporary tyre after the original forest has been
burned-over or otherwise denuded. The Larix type is believed
to be a temporary type prior to the Swamp Conifers. The
Swamp Hardwoods, the Swamp Conifers of Thgjaegbiegfgigeg,

and the Marsh type are all to be the subclimax type to the
region.

Silvicultural treatments are considered for the three
forest types, the Swamp Hardwoods, the Swamp Conifers, and
the lowland ASpen type. Light partial cutting is believed
to be a good measure for the Swamp Hardwood type. Clear cutt-
ing in strips is suggested for the Swamp Conifer type. Clear
cutting is probably the only method for the Lowland Aspen

type.

APPENDIX A

List of common names of plants used with scientific equivalents
Alder, green ---- Alngg sinuats (Reg.) Rydb.
Alder, speckled ---- Alggg rugosa (Du Roi) Spreng.
Ash, black ---- Fraxinus giggg Marsh.
Ash, green --—- Fraxinus pgnpsylvanica Marsh.
Aspen, trembling ---- Porulus tremuloides Michx.
Basswood -—-- Tilig americana L.
Birch, swamp ---- Betula pumila L. .
Birch, sweet ---— Betula lenta L.
Birch, yellow —--- Betula lutea Michx, f.
Bluejoint —--- Calamagrostis canadengig (Michx.) Nutt.
Cat-tail ---- Typha latifolia L.
Cedar, northern white ---- Thuja occidentglig L.
Cherry, black -—-- Eggnus gerotigg Ehrh.
Cherry, choke ---— Prunug Virginiana L.
Cherry, fire ---- Prunus.pensylvanica L. f.
Dogwood, red osier -~-- Cornus stolonifera Michx.
Elm, American ---- glmus americagg L.
Elm, rock --—- Elma; thomasi; Sarg.
Elm, slippery ---- glmgg ggpgg Muhl.
Fern, royal ---- Osmunda regalis L.
Fir, balsam ---- Abies balsamea (L.) Mill.
Hemlock -—-- Tsuga canadensis (L.) Carr.

Hornbeam, American -—-- Carpinus caroliniana Walt.

170

Maple, red ---- Acer rubrum L.

Maple, silver
Oak, northern
Pine, eastern
Spruce, black
Tamarack ----

Tea, Labrador

Willow, heart-

 

---- Acer saccharinum L.

red ---— Quercus rubra L.

white ---- Einus Strobus L.
---- Picea mariana (Mill.) ESP.
larix laricina (Du Roi) K. Koch
---- Ledgg groenlandicum Oeder.

leaf ---- Salix eriocephala Michx.

Willow, long—beaked---- Salix bebbiana Sarg.

171
APPENDIX B
List of plants encountered in making the vegetation

study in the Higgins Lake area.

TREES

 

1.A§ig§'balsamea (L.) M111.
2.Acer rubrum L.
B.Agggygaccharinum L.
4.Betula lgngg L.

*fi5.Betula lutg§,Michx. f.
6.Ca;pinus caroliniana Walt.
7.Fraxinus,gig§g,Marsh.

*8.Fraxinus pennsylyanica Marsh.
9.Lg§i; laricina (Du Roi) K. Koch
10.2iggg mariana (Mill.) ESP.
11.21nus Strobus L.

l2.P0pulus tremuloides Mickx.
13.Prunus serotina Ehrh.

*14.Qg§rcus rubra L.

15.2hglg occidentalis L.
16.1111; americang L.
l7.Tsuga canadensis (L.) Carr.
18.leus amerigana L.
19.leg§ thomasii Sarg.
20.Ulmus rubra Muhl.
**Betu1a alleghaniensig'Britton in ”Check List of Native
and Naturized Trees of the United States (Including

Alska)‘
*After the "Check List" (U.S.D.A.F.S. Handbook No.41,1953)

ggggss AND vgggg
1.Acer spicatum Lam.
*2.§;gg§ ginuata (Reg.) Rydb.
3._;ng§ rugosa (Du Roi) Spreng.
4.Amelanch1er arborea (Michx. f.) Fern.

B.Amelanchier intermedia Spach

 

6.Amelanchier sp.
*7.Aronia nigra.Britt.
B.Asclepias incarnata L.

9.Betu;a pgmila L.
10.Clematis virginiana L.
11.Cornus alternifolia L. f.
l2.Co;nus gtolonifera Michx.
13. gulps cornuta Marsh.

14.G§gthgria procumbens L.
15.Ilex verticillata (L.) Gray

16.Ledum groenlandicum Oeder.
17.L1nnaea borealis var. gmericana (Forbes) Rehd.

 

 

 

18.Lonicera canadensis Bartr.

19.Lonicera oblongifolia (Goldie) Hook.

20.Lonicera villosa var. solonis (Eat.) Fern.

21.Mitche11a repens L.
22.Nemopanthus mucronata (L.) Trel.

23. agthenocissus inserts (Kerner) K. Fritsch.
24.Poten§111a fructicosa L.

 

 

25.Prunus ayium L.
*After "Check List".

‘

172

26.Prunus pensylvanica L. f.

27.Prunus Virginiana L.

28.3Qus radicans L.

29.Ribes

30.Ribes

31.Ribes
32.3222;
33.3gpgg
34.Rubus
35.3222a
36.3222a
37-.éll£
38.§alie
39.§all£
40.§§11;
*4l.§aliz
42.§g;;g.
*4}.§a11£
44-éalia
45.§allz

émericana Mill.

lacustre (Pers.) Poir.

trieste Pall.

flagellaris Willd.

hiaaidaa L-

ideaus L.

ideaus var. aculeatissimus Regel & Tiling
pubescens Raf.

alba var. vitellina (L4) Stokes
amygdaloides Anderss.

bebbiana Sarg.

discolor Mflhl.

eriocephala Michx.

lucida Muhl.

petiolaris J.E.Sm.

gerissima (Bailey) Fern.

Sp.

46.Smilax tamnoides var. hispida (Muh1.) Fern.

*47.Sorbus americana Marsh.

48.321raea alba Du Roi

49.Spiraea latifolia (Ait.) Borkh.

50.Vaccinium myrtilloides Michx.

51.Vaccinium vacillans Torr.

*After "Check-List".

173

174

PTERIDOPHYTES
1.Botrychium dissectum Spreng.
2.Cystopteris bulbifera (L.) Bernh.
3.Dry0pteris cristata (L.) Gray
4.Dr10pteris cristata var. clintoniana (D.C.Eat.) Underw.
5.Dryopteris spinulosa (O.F. Muell.) Watt.
6.Dryopteris thelypteris (L.) Gray
7.§ggisetum Lalustre L.
8.§ggisetum sylvaticum L.
9.0nac1ea sensibilis L.

10.0smunda regalis L.

SEUGES, GRASSES,7AKD HERBS
l.Agropyron repens (L.) Beauv.
2.Agrostis hyemalis (Walt.) Beauv.
3.Bromus ciliatus L.
4.Bromus purgans L.
5.0a1amagrostis canadensis (Michx.) Mutt.
6.Carex comosa Boott.
7.Carex crinita Lam.
8.Carex diandra Schrank
9.Carex intumescens Rudge
10.9aggz,lasiocarpa Ehrh.
11.Qa§§1 tenera Dew.
12.Carex trispermg Dew.
13.Eleocharis obtusa (Willd.) Schultes
l4.Glyceria boreglig (Nash) Ratchelder
15.Glyceria canadensis (Michx.) Trin.
16.Glyceria pallida (Torr.) Trin.
17.glygeria striata (Lam.) Hitchc.
18.ggpcus canadensis J. Gay

19.Muhlenbergia racemosa (Michx.) ESP.

20.Scirpus cyperinus (L.) Kunth

21.Alisma triviale Pursh.
22.Aralia nudicaulis L.
23.§§tg§¢1gnciformis'Rydb.
24.A§tgg lateriflorus (L.) Britt.

25.Agter novae-angliae L.

26.A§ter puniceus L.

27 .Agtgg sp.

28.Bidens cernua L.

29.§idens frondgga L.

30.0ampanula aparinoideg Pursh.
31.91guta bulbifera L.

32.Circium muticum Michx.
33.9;gntonia borealis; (Ait.) Raf.
34.Coptis greenlandica (Oeder) Fern.

35.Cornus canadensis L.

36.§pilobium glandglosum var. adenocaulon (Haussk.) Fern.

37.Epilobium leptonhyllum Raf.
38.Eupatorium perfoliatum L.
39.Eupatorium purpureum L.
40.E§agaria vesca L.

41.Fragaria virggniana Duchesne.

42.Fragaria Virginians var. illinoensis Gray

 

43.Galium aparine L.
44.9glium asprellum Michx.
35.Galium obtusum Bigel.
46.Galium tinctorium L.
47.Galium triflorum Michx.
48.Galium trgfidum L.

49.Galium ep.

176

177

50.Geum aleppicum var. strictum (Ait.) Fern.
51.Goodyear pubescens (Willd.) R. Br.

52.Habenaria obtusata (Pursh.) Richards.

 

53.Hypericum boreale (Britt.) Bickn.
54.;mpatiens capengig Meerb.
55.;gpatiens pallida Nutt.

56.;gis versicolor L.

57.Lemna minor L.

58.L§mna trisulca L.

59.Lycopus americanus Muhl.
60.;gpopus rubellus Moench.
61.Lysimachia nummularia L.
62.Lysimachia terrestris (L.) ESP.
63.1gsimachia thyrsiflora L.
64.Maianthemum canadensis Desf.
65.Mentha arvensis L.

66.Mite11a nuda L.

67.2;pnellg vulgaris L.

68.Pyrola rotundifolia (L.) Desf.
69.2yrola ggcundg L.

70.Rannucu1us sp.

71.3gmex verticillatus L.

*72.Sagittaria montevidensis Cham. & Schlecht.
73.§ggtellaria epilobiifolia A. Hamilton
74.8cute11aria 1gpepiflora L. I

“After Gleason,H.A. 1952. New Britton & Brown Illustrated

Flora. The New York Botanical Garden.
Lophotocarpus calycinus in Gray's Manual of Botany.

178
75.31um suave Walt.
76.5m11gcipg trifolia (L.) Desf.
77.Solidago caesia L.
78.Solidago canadensig L.
79.Solidago graminifolia (L.) Salisb.
80.§glidago missougiensis Nutt.
81.§g;;dago patula Muhl.
82.Solidago rugosa Ait.
83.Solidago ulmifolia Muhl.
84.Sol;dago uliginosa Mutt.
85.3pirode1a polyrhiza (L.) Schleid.
86.Thalictrum dasycarpum Fisch. & Lall.
87.Trientalis borealis Raf.
88.Typha latifolia L.
89.Veronica ggutellaria L.

90.!i91a blanda Willd.

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190

 

 

Plate 3. A Scirpgs-Txpha stand of the Marsh type. The
dominance of cat-tails (Typha latifolia) is indicated
by their numerous flowering stems in the fore-ground.
An Aspen (Pogglus tremuloides) stand is in the back
ground. In between, willows (Salix sp.) are grown in
low bushes to give a patchy apperance.

 

.s . It." .m
lt-O, it“s,eljrt t

191

 

Plate #. A Sglix-Calam roetis stand of the Marsh
type. An Aspen (Popglus tremuloides) stand is
in the back ground.

192

 

Plate 5. Soil profile of the Newton loamy sand in a
Salix—Calamagrostis stand of the Marsh type during
the dry season in October. The water-table was
down to a depth of approximately 2 feet. The organic
peat layer reached down to about 1% feet.

193

 

A typical view of the Swamp Coniferous Forest.

Plate 6.

194

 

Plate 7. External apperance of the Elm-Soft Maple stand
of the Swamp Hardwoods type.

 

Plate 8. Ground vegetation of the Elm-Soft Maple stand
of the Swamp Hardwoods type showing seedlings of maple,
elm, and ash. Carex intumescens can be seen in the
fore-ground with the inflorescences shown in center

of the picture.

 

195

 

Soil profile of the Bergland loam in the
Wateritable was at approximately 10 inches.

Ash-Red Maple-Elm stand of the Swamp Hardwoods

“Plate 9.
type.

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