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An investigation of the periodicity of Gelidium cartilagineum, a perennial red alga

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AN INVESTIGATION OF THE PERIODICITY OF GELIDIUM
CARTILAGINEUM, A PERENNIAL RED ALGA
A Thesis
Presented to
the Faculty of the Department of Botany
University of Southern California
In Partial Fulfillment
of the Requirements for the Degree
Master of Arts
by
Fenton Lawrence Feeney
February 1940
UMI Number: EP41408
All rights reserved
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UMI EP41408
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presented to and accepted by the C o u n c i l on
G r a d u a t e S t u d y a n d Research in p a r t i a l f u l f i l l ­
m e n t o f t h e r e q u i r e m e n t s f o r th e d e g r e e o f
MASTER OF ARTS
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r....... ...........
-
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Secretary
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February.. 19 40....
F aculty Com m ittee
Chairm an
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ACKNOWLEDGMENTS
To Dr. George R. Johnstone, the writer wishes to
express his sincere gratitude for his guidance through­
out the research and the writing of this paper.
Appreciation is also accorded to Ifr. S.H. Corfield
of the United States Agar Company in San Diego, California
for his cooperation in supplying some of the material for
the investigation.
Expressions of appreciation are also due friends
who rendered their services: Mr. Lester Barcus, photo­
graphy; Mr. H.W.G. Bertling, translation; and Mr. Glenn
L. Arbogast, Jr., and Mr. Peter K. Ballou, members of the
Explorer Patrol, Boy Scouts of America, for aid on col­
lections.
TABLE OF CONTENTS
PAGE
I N T R O DUCTION.........................................
1
The S p e c i e s ............ . . .. ...................
1
Statement of the Problem ..........................
2
Justification of the Problem • • . • • • • • • • •
2
TECHNIQUE AND P R O C E D U R E ............................
Sources of Material
4
.. . . .......................
4
................................
5
RESULTS AND DISCUSSION ..............................
8
SUMMARY AND C O N C L U S I O N S ............................
22
LITERATURE C I T E D ................................ ..
23
Method . . . . . .
.
A P P E N D I X .............................................
24
LIST OF FIGURES
FIGURE
1
2
3
PAGE
The vegetative structure of the three indivi­
duals are morphologically similar and macroscopically indistinguishable, above - 0,
female;
®, tetrasporic; and 0, male plant.
The microscope reveals, below, three types
of individuals: a. antheridial; b. cystocarpic, and c. tetrasporic.
From unpublish­
ed report of Johnstone and Wells (1934) . . .
7
Most of a mature plant showing the new basal
shoots in relation to the adult shoots
• • •
17
New basal shoots as they appear on the
stoloniferous holdfast.
This is an
enlargement of a lower portion of Figure
2 ....................................
19
LIST OF TABLES
TABLE
I.
II.
III.
IV.
V.
VI.
VII.
PAGE
GELIDIUM CARTILAGINEUM COLLECTED AT LAGUNA
BEACH, CALIFORNIA TABULATED FROM APPENDIX
TABLE VII TO SHOW THE NUMBER AND PERCENTAGE
FREQUENCY OF THE VARIOUS TYPES OF INDIVI­
DUALS: ANTHERIDIAL, CYSTOCARPIC, AND TETRA­
SPORIC ......................................
9
THE PERCENTAGE FREQUENCIES OF ANTHERIDIAL,
CYSTOCARPIC, AND TETRASPORIC PLANTS COLLECT­
ED AT WHITE POINT FROM JANUARY 1935 TO JUNE
1934 EXCEPTING THE COLLECTION OF MARCH 11,
1954 FROM THE BEACH 3 MILES NORTHWEST OF
TOPANGA CANYON AND THAT FOR APRIL 8, 1934
FROM PORTUGUESE BEND ABOUT 2 MILES WEST OF
SAN PEDRO, CALIFORNIA
......... . .........
10
PERCENTAGE FREQUENCY OF ANTHERIDIAL, CYSTO­
CARPIC, TETRASPORIC PLANTS, AND UNDETER­
MINED FORMS OF 100 SPECIMENS TAKEN FROM
LARGE COLLECTIONS MADE ABOUT THE FIRST AND
THIRD WEEKS IN JUNE, 1939. MATERIAL FROM
MR. CORFIELD OF THE U.S. AGAR COMPANY, SAN
DIEGO, CALIFORNIA. COLLECTED ALONG THE SAN
PEDRO BREAKWATER - ABOUT FIFTEEN FEET BELOW
THE SURFACE OF THE WATER ...................
12
THE FREQUENCY OF UNDETERMINED SPECIMENS WHICH
SHOW INJURED BRANCHES AND THE DEGREE OF RE­
GENERATION THAT HAS TAKEN PLACE. TABULATED
FROM APPENDIX TABLE VIII ...................
14
THE FREQUENCY OF VEGETATIVE ACTIVITY AMONG 10
ADULT SHOOTS AND THE FREQUENCY OF OCCURRENCE
OF NEW BASAL SHOOTS FOR 10 SPECIMENS OF EACH
COLLECTION.
TABULATED FROM APPENDIX TABLE
I X ...........................................
15
THE NUMERICAL RELATION AND RATIO OF TOTAL
FREQUENCY OF ADULT SHOOTS TO NEW SHOOTS
FOR EACH COLLECTION.
TABULATED FROM APPEN­
DIX TABLE X
................................
21
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULAT­
ED TO SHOW THE NUMBER AND PERCENTAGE FRE­
QUENCY OF THE VARIOUS TYPES OF INDIVIDUALS .
24
LIST OF TABLES, CONTINUED
TABLE
VIII.
PAGE
THE EXTENT OF INJURED BRANCHES AND THE
DEGREE OF REGENERATION AMONG SPECIMENS
UNDETERMINED AS TO TYPES OF REPRODUCTIVE
A C T I V I T Y .......................................
IX.
DEGREE OF ACTIVITY AMONG ADULT SHOOTS (x),
AND THE OCCURRENCE OF NEW BASAL SHOOTS
(x)
X.
THE NUMERICAL RELATION OF FREQUENCY OF
ADULT SHOOTS (NUMERATOR) TO NEW SHOOTS
(DENOMINATOR) AMONG INDIVIDUAL HERBARIUM
S P E C I M E N S ...........................
32
5
INTRODUCTION
The rhythmic succession of vegetative and repro­
ductive activities is a usual phenomenon and apparently a
necessity for survival among terrestrial plants in habitats
where marked seasonal changes in temperature, moisture, and
light prevail.^
A similar sequence of activities is found
among annual aquatic plants, but perennial marine plants
along the coast of southern California present a less pro­
nounced periodicity of growth and reproduction.
In such
an environment seasonal changes are greatly reduced compar­
ed to those which prevail in adjacent terrestrial habitats.
Gelidium cartilagineum is a perennial red alga which grows
at and below the lowest tide level where changes in tempera­
ture and light are much reduced and desiccation never occurs.
The Species. - By comparing the anatomical work done
in the laboratory with the anatomical descriptions and il­
lustrations of Kylin (1928), the species investigated in
this study was determined as Gelidium cartilagineum.
It
appears to be G. cartilagineum (1.) Gaill. as described by
Kylin (1925, p. 15).
A specimen from Ensenada, Lower
California, was described and named G. cartilagineum var.
robustum var. nov. by Gardner (1927, p. 280) who states,
"I have not seen the specimen reported by Kylin (1925, p.
13) from San Juan Island, ¥/ashington, but probably it be-
•^A good discussion of growth periodicity of higher
plants may be found in Meyer and Anderson, Plant Physiology.
1939.
2
longs here*
If so the range of the stated distribution is
to be extended.”
Statement of the Problem. - Morphological alterna­
tion of generations occurs in the life cycle of Gelidium
cartilagineum and Its gametophyte generation Is dioecious
(Kylin, 1928).
The vegetative structures of the three in­
dividuals, tetrasporic, antheridial, and cystocarpic plants
are morphologically similar and macroscopically indistin­
guishable.
Any evidence of periodicity in the vegetative
structures must be determined by a study of new growth at
the bases and apices together with regeneration of injured
branches, while an attempt to determine periodicity in re­
productive activities may be made by noting the frequency
of occurrence of reproductive structures on a percentile
basis.
Such evidence can be obtained among specimens which
have been collected at regular monthly intervals.
Justification of the Problem, - Gelidium cartilagineum
has been collected off the coast of southern California for
many years for the purpose of manufacturing agar-agar of com­
merce.
In 1930 Mr. H.D. McKinnon of the American Agar Com­
pany in San Diego raised the question as to whether or not
this marine plant might not reach extinction due to the con­
tinuous harvesting throughout the year.
A knowledge of the
periodicity of this marine plant should prove helpful in
ansveering such a question.
Furthermore, the use of agar-
agar is becoming more prominent in the United States and
Europe.
Tressler (1923) states that agar-agar has long been
3
used for food in the Orient*
He explains that it is eaten
in the form of jellies and as a thickener of soups, sauces,
and gravies.
It is also eaten in the United States and
Europe chiefly in food preparations such as ice cream,
jellies, candies, pastries and in many desserts.
Its chief
uses in Europe and America are as a colloidal matrix in pre­
paration of nutrient media for the culture of bacteria and
molds and in medicine for treatment of chronic constipation.
Although this problem has a popular or commercial aspect,
its significance in building up our knowledge of periodicity
in perennial marine plants should not be overlooked.
TECHNIQUE AND PROCEDURE
The sources of material and methods of investigation
are presented in this section.
Sources of Material. - Since many of the red algae
including Gelidium cartilagineum grow at some depth below
the surface of the water this species was gathered at the
second lowest tide of the month, usually every fourth week.
Of the fifteen trips taken from February 13, 1938 to April
1, 1939 there are no data for three trips owing to the fact
that even the lowest tide of the month was too high or the
surf too severe for collecting.
One series of collections
was done at Laguna Beach, California except for the month
of March 1938.
The March collection was made at Point
Firmin, California, because of the road conditions along
the coast between Seal Beach and Laguna Beach after the
flood of March 2.
The temperature of the water was record­
ed for each collection beginning in May 1938.
The collect­
ed material was pressed and dried in a plant press.
The aim
was to get one hundred specimens of separate individual
plants for each collection.
However, due to the conditions
provided by the tides, the number of specimens among the col­
lections range from fifty-one to one-hundred-five.
The data on the above material collected at Laguna
Beach and Point Firmin, California have been supplemented
with data on similar collections obtained near the Break­
water at Cabrillo Beach, San Pedro, California through the
5
courtesy of Mr. S.H. Corfield of the United States Agar
Company of San Diego, California.
Other data on material
collected at White Point, about two miles west of San Pedro,
California have been added through the courtesy of Dr. G.R.
Johnstone and Leta Wells.
Method. - A part of each specimen was examined under
the microscope to determine its type: antheridial, cysto­
carpic, or tetrasporic (Figure 1, a b c), or where sufficient
differentiation was lacking the specimen was designated as
undetermined.
These undetermined specimens which did not
show reproductive characteristics were examined fof indivi­
duals with injured branches and for regeneration.
Ten specimens from each of the Laguna Beach collec­
tions were also selected because of their apical and basal
vegetative activity.
These ten representatives of each
collection were compared with the representatives of all
other Laguna Beach collections to determine whether or not
periodicity occurs in the vegetative activity.
Figure 1
The vegetative structures of the
three individuals are morphologi­
cally similar and macroscopically
indistinguishable, above —
(j),
female; ©* tetrasporic; and C?,
male plant.
The microscope reveals
below, three types of individuals:
a, antheridial, b. cystocarpic, and
c. tetrasporic.
From unpublished
report of Johnstone and Wells (1934).
-
22
-
RESULTS AND DISCUSSION
Data on reproductive and vegetative activities are
presented from examinations of the three sets of material.
The collection from Laguna Beach, California showed consid­
erable injury and severing of branches due to pounding of
the surf.
This material, however, proved to be of great
value in the study of regeneration for which data are also
presented.
A comparison of the data on percentage frequencies
of the male and female plants together with the tetrasporic
plants in the columns on reproduction in Table I, shows the
greatest percentage frequency of tetrasporic individuals
for all months except July 1938.
The column of undeter­
mined specimens was necessarily omitted in this comparison.
These undetermined specimens were so badly injured by the
surf that they could not be classified among the three
types of reproduction.
The number of undetermined speci­
mens and the collecting conditions result in data that are
not truly representative of reproductive activity for cer­
tain months.
However, the predominance of tetrasporic
plants indicates that asexual reproduction is more frequent
than sexual.
Data from material collected at White Point
and determined by G.R. Johnstone and Leta Wells in 1933 and
1934 (Table II) are more consistant than the data in Table
I, indicating without exception the largest percentage fre­
quency for tetrasporic individuals.
These data are more
9
TABLE I
GELIDIUM CARTILAGINEUM COLLECTED AT LAGUNA BEACH, CALIFORNIA
TABULATED FROM APPENDIX TABLEVII TO SHOW THE NUMBER AND PER­
CENTAGE FREQUENCY OF THE VARIOUS TYPES OF INDIVIDUALS: ANTHIDIAL, CYSTOCARPIC, AND TETRASPORIC
Antheridial Cystocarpic Tetrasporic Undetermined
Date:
No. per
No. per
Yr. Mo. Da. No. per
No. per
cent
cent
cent
cent
16 18.3
33 37.9
18 20.6
1938 2 13 20 22.9
14.8
13
26
12.9
8
48.1
7
13 24.0
*1938 3
3.8
58 55.7
0.96
4
1
41 39.4
1938 4 10
8 11 12.9
8.2
56 65.8
7
1938 5
11 12.9
3
1938 6
33 64.7
1.9
3 17 33.3
1
1938 7
23 34.3
19 28.2
2.9
2
23 34.3
1938 7 30
1938 8 28
4
8 12.6
6.3
25 39.6
26 41.2
1938 9 25
14 21.2
3.03
10 15.1
2
40 60.6
1938 10 23
8
9.3
2.3
25 29.06
2
51 59.3
1938 11 20
1938 12 19
14 15.5
26 28.8
30 33.3
2 20 22.2
1939 2
9 14.7
3
4.9
49 80.3
5
1939 3
14 13.3
23 21.9
7.6
60 57.1
8
1
1939 4
*This material was collected at Point Firmin because of road
conditions after the flood of March 2, 1938.
10
TABLE II
THE. PERCENTAGE FREQUENCIES OF ANTHERIDIAL, CYSTOCARPIC,
AND TETRASPORIC PLANTS COLLECTED AT VtfHITE POINT FROM
JANUARY 1953 TO JUNE 1934 EXCEPTING THE COLLECTION OF
MARCH 11, 1934 FROM THE BEACH 3 MILES NORTHWEST OF
TOPANGA CANYON AND THAT FOR APRIL 8, 1934 FROM PORTU­
GUESE BEND ABOUT 2 MILES WEST OF SAN PEDRO, CALIFORNIA
Date
1933
Jam
Feb,
Mar.
Apr.
May
June
July
Aug.
Sept.
Oct.
Nov.
Dee.
Antheridial
Cystocarpic
Tetrasporic
23
22
22
13
12
11
9
6
23
20
17
16
18
0
16
23
11
10
1
4
0
42
0
10
26
3
1
8
3
0
7
7
15
1
17
0
56
97
83
69
86
90
92
89
85
57
83
90
16
11
11
12
8
9
4
1
17
9
10
4
13
13
21
17
14
8
3
8
0
1
0
11
69
83
87
88
87
86
79
72
1934
Jan.
Feb.
Mar.
Apr.
Apr.
Apr.
May
June
truly representative of reproductive activities for the mat
erial from thich these data were obtained showed no injured
forms.
Additional data on injured specimens are presented
in Table III which show the percentage frequency of antheri
dial, cystocarpic, tetrasporic plants, and undetermined
forms of 100 specimens collected along the San Pedro break­
water on two different occasions also indicates that tetra­
sporic individuals occur In the greatest percentage frequen
cy.
This San Pedro material was collected at a level of
about fifteen feet below the surface of the water where in­
jury from the surf was impossible, while the Laguna Beach
and White Point material were collected along the shore at
low tide.
With the frequency of tetrasporic individuals
so similar in each of the localities, it can be said that
the influence of habitat has little or no effect upon the
percentage frequencies with the exception of the indirect
effect due to injury caused by the surf at low tide.
Haupt (1932) states in his work on Zonaria farlowii
that three kinds of individuals superficially alike are
present, viz., male, female, and asexual, and that the
sexual plants are comparatively rare.
Robinson (1932)
also reported a rarity of the sexual plants of Toania.
These data on Zonaria and Toania, brown algae of the family
Dictyotaceae and on Gelidium. a perennial red alga, support
the conclusions that there is inhibition of sexual plants
and thus a greater preponderance of non-sexual plants.
The surf at low tide injured the undetermined speci-
12
TABLE III
PERCENTAGE FREQUENCY OF ANTHERIDIAL, CYSTOCARPIC, TETRA­
SPORIC PLANTS, AND UNDETERMINED FORMS OF 100 SPECIMENS
TAKEN FROM LARGE COLLECTIONS MADE ABOUT THE FIRST AND
THIRD WEEKS IN JUNE, 1939.
MATERIAL FROM MR. CORFIELD
OF THE U.S. AGAR COMPANY, SAN DIEGO, CALIFORNIA. COLLECT­
ED ALONG THE SAN PEDRO BREAKWATER - ABOUT FIFTEEN FEET
BELOW THE SURFACE OF THE WATER
Week
Antheridial
Cystocarpic
Tetrasporic
Undetermined
1
7
10
76
7
3
8
IS
77
3
13
mens to the extent that the nature of their reproductive
structures was beyond determination but these specimens
proved to be of great value in a subsequent study of re­
generation.
Table IV presents a tabulation of the fre­
quency of undetermined specimens showing Injured branches
and the degree of regeneration, and shows a predominance
of regeneration in injured material collected during the
spring and fall months.
These data are in agreement with
the observations of Svedelius (1906) who states that period­
icity appears in perennial forms with strong basal parts,
often after fructification, by dropping old shoots and re­
placing them by new ones.
The examples cited by him were
Laurencia cevlanica and Rhodomela crassicaulisf red algae;
Sargassum cristaefoliumf a brown alga, and Aurainvillea
lacerata. a green alga.
Table V shows the frequency of vegetative activity
among adult shoots and the frequency of occurrence of new
basal shoots for each collection.
Each month shows some
vegetative activity among the adult shoots.
February and
March may be designated as the early peripd for most of the
vegetative activity and October as the later period.
New
basal shoots (Figures 2 and 3) appear from stoloniferous
holdfasts (Figure 3) every month in the year.
These basal
shoots vary from three to four inches in length before
branching begins.
The number of adult shoots is compared with the num-
14
TABLE IV
THE FREQUENCY OF UNDETERMINED SPECIMENS WHICH SHOW INJURED
BRANCHES AND THE DEGREE OF REGENERATION THAT HAS TAKEN
PLACE. TABULATED FROM APPENDIX TABLE VIII
Date
Number
observed
Injured:
Slightly
Severely
Not
Regeneration •
No Fair
Good
1938
2-13
3-13
4-29
5-8
7-3
7-30
9-25
10-23
11-20
18
11
38
55
26
23
23
36
45
9
3
10
18
10
6
10
10
17
6
4
27
20
8
8
9
24
23
3
4
1
17
8
9
4
2
5
4
3
21
18
18
18
15
21
22
14
5
14
35
7
5
7
13
20
0
3
3
2
1
0
1
2
3
14
40
61
4
11
14
6
29
42
4
0
5
11
21
22
2
17
34
1
2
5
1939
2-2
3-5
4-1
15
TABLE V
THE FREQUENCY OF VEGETATIVE ACTIVITY AMONG 10 ADULT SHOOTS
AND THE FREQUENCY OF OCCURRENCE OF NEW BASAL SHOOTS FOR 10
SPECIMENS OF EACH COLLECTION.
TABULATED FROM APPENDIX
TABLE IX
Date
Apical activity •
Good
Fair
Slight
Basal activity:
Presence only
1958
2-15
5-13
4-10
5-8
7-3
7-30
9-25
10-23
11-20
4
0
0
0
4
0
1
7
3
4
3
6
7
4
5
7
2
6
2
7
4
3
2
5
2
1
1
10
10
7
9
10
9
10
10
10
2
7
2
6
3
6
2
0
2
10
10
9
1939
2-2
3-5
4-1
Figure 2
Most of a mature plant show­
ing the new basal shoots in
relation to the adult shoots.
About natural size.
Figure 3
New basal shoots as they appear
on the stoloniferous holdfast*
This is an enlargement of a low­
er portion of Figure 2.
Enlarged five times.
•V
'•
20
ber of new shoots in Table VI.
New shoots are those basal
shoots that show no lateral branching, while adult shoots
are those with lateral branches.
The greater number of
new shoots in comparison to adult shoots occurs during the
months of March and July, which indicates the greater veg­
etative activity during these months.
The ratio of new
shoots to adult shoots is less than two for the other
months of the year.
21
TABLE VI
THE NUMERICAL RELATION AND RATIO OF TOTAL FREQUENCY OF
ADULT SHOOTS TO NEW SHOOTS FOR EACH COLLECTION
TABULATED FROM APPENDIX TABLE X
Date
1938
Adult
2-13
3-13
4-10
5-8
7-3
7-30
9-25
10-23
11-20
117
78
74
39
103
44
94
121
87
87
33
42
32
51
48
93
82
65
1.34
2.36
1.76
1.21
2.01
0.91
1.01
1.47
1.33
78
122
99
43
70
51
1.81
1.74
1.94
New
Ratio A/N*
1939
2-2
3-5
4-1
*A - Adult shoots; N - New shoots
SUMMARY AND CONCLUSIONS
1.
An investigation of the vegetative and repro­
ductive structures of Gelidium cartilagineum. a perennial
red alga, was made of material collected on fifteen monthly
trips to Laguna Beach, California together with additional
material from Cabrillo Beach and White Point near San Pedro,
California.
2.
The three morphologically similar types of indi­
viduals which appear in the life cycle of this species dif­
fer in their frequencies of occurrence.
The tetrasporic
plant occurs more frequently than either the male or female
plant indicating that asexual reproduction is more frequent
than sexual reproduction.
3.
Regeneration of injured vegetative parts of adult
shoots occurs chiefly during the spring and fall months.
4.
Data are presented which indicate that the adult
shoots show two periods of vegetative activity, an early per­
iod of February and March and a late period in October.
5.
New basal shoots show no periodicity.
They vary
from one to four inches in length and these variations occur
every month in the year.
6.
Results indicate that the time of harvest of
Gelidium cartilagineum for commercial uses is not of parti­
cular importance so far as extermination of the species is
concerned in the areas from which material for study had
been collected.
LITERATURE CITED
Gardner, N.L., New species of Gelidium on the Pacific
1927
Coast of North America,
Univ, Calif, Publ. Bot., 13:273-318.
Raupt, Arthur W,, Structure and development of Zonaria
1932
farlowii,
Johnstone, George R,, and Leta Wells, Observations of
1934
Gelidium cartilagineum var, robustum.
Unpublished report,
Kylin, Harold, Entwicklungsgeschichliche Floridienstudien
1928
Lunds. Univ, Arssk, N.F. Avd, 2 Bd. 24
Nr 4
Kylin, Harold, The marine algae in the vicinity of the
1925
biological station at Friday Harbor, Wash*
Lunds, Univ, Arssk, N.F. Avd. 2 Bd. 24
Nr 4
Meyer, Bernard S., and Donald B. Anderson, Plant Physiology.
1939
D. van Nostrand Company, Inc.
250 Fourth Avenue, New York City
Svedelius, Nils, Uber die Algenvegetation eines ceylonischen
1906
Korallenriffes mit besonderer R-ftchsicht auf
ihre Periodizit&t.
Botaniska Studier. Uppsala, Sweden
Tressler, Donald K., Marine Products of Commerce
1923
Chemical Catalog Company, Inc.,
New York.
24
TABLE VII
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of February 13, 1938, Laguna Beach, Californis
Tide: 2:41 P.M.
-1.4
Temperature (water): ---Number
Percent
Antheridial
20
22 •9
Cystocarpic
16
18.3
Tetrasporic
33
37.9
Undetermined
18
20.6
Total
87
Collection of March 13, 1938, Point Firmin, California
Tide: 1.37 P.M.
-0.9
Temperature (water): ---Number
Percent
Antheridial
7
12.9
Cystocarpic
8
14.8
Tetrasporic
26
H
»
CO
Undetermined
13
24.0
Total
54
25
TABLE VII, CONTINUED
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of April 10, 1938, Laguna Beach, California
Tide: 12.26 P.M.
-0.3
Temperature (water): ---Number
Percent
Antheridial
1
.96
Cystocarpic
4
3.84
Tetrasporic
58
55.7
Undetermined
41
39.42
Total
104
Collection of May 8, 1958, Laguna Beach, California
Tide: 11.00 A.M.
0.2
Temperature (water): 19° C.
Number
Percent
Antheridial
11
12.9
Cystocarpic
7
8.2
Tetrasporic
11
12.9
Undetermined
56
65.8
Total
85
26
TABLE VII, CONTINUED
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of June 3, 1938, Laguna Beach, California
Tide: 7:21 A.M.
-0*4
Temperature (water): 17*5° C.
Number
Percent
Antheridial
Cystocarpic
Collecting Poor
No Material
Tetrasporic
Undetermined
Collection of July 3, 1938, Laguna Beach, California
Tide: 7*28 A ♦M.
0.6
Temperature (water): 18.5° C.
Number
Percent
Antheridial
17
o3 *3
Cystocarpic
0
0*0
Tetrasporic
1
1.9
33
64.7
Undetermined
Total
51
27
TABLE VI, CONTINUED
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of July 30, 1938, Laguna Beach, California
Tide: 5:15 A.M.
-0.1
Temperature (water): 18.5° C.
Number
Percent
Antheridial
2
2.9
Cystocarpic
23
34.3
Tetrasporic
19
28.2
Undetermined
23
34.3
Total
67
Collection of August 28, 1938, Laguna Beach, California
Tide: 5:14 P.M.
0.7
Temperature (water): 22° C.
Number
Percent
Antheridial
Cystocarpic
Tetrasporic
Undetermined
Collecting Poor
No Material
28
TABLE VII, CONTINUED
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of September 25, 1938, Laguna Beach, California
Tide: 4:06 P.M.
0.0
Temperature (water): 20° C.
Number
Percent
Antheridial
4
6.3
Cystocarpic
8
12.6
Tetrasporic
25
39.6
Undetermined
26
41.2
63
Total
Collection of October 23, 1938, Laguna Beach, California
Tide: 3:13 P.M.
-0.5
Temperature (water): 18.0° c.
Number
Percent
Antheridial
2
3.03
Cystocarpic
10
15.10
Tetrasporic
14
21.20
Undetermined
40
60.60
Total
66
29
TABLE VII, CONTINUED
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of November 20, 1938, Laguna Beach, California
Tide: 2:30 P.M.
-0.6
Temperature (water): 17° C.
Number
Percent
Antheridial
2
2.30
Cystocarpic
8
9.30
Tetrasporic
25
29.06
Undetermined
53
59.20
-
Total
86
Collection of December 19, 1938, Laguna Beach, California
Tide: 2:25 P.M.
-0.7
Temperature (water): 16.5° C.
Number
Percent
Antheridial
Cystocarpic
Tetrasporic
Undetermined
Collecting Poor
No Material
50
TABLE VII, CONTINUED
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of February 2, 1959, Laguna Beach, California
Tide: 2:15 P.M.
-1.5
Temperature (water): 13.5 ° C.
Number
Percent
Antheridial
20
22.2
Cystocarpic
26
28.8
Tetrasporic
30
33.3
Undetermined
14
15.5
Total
90
Collection of March 5, 1939 , Laguna Beach, California
Tide: 3:03 P.M.
-1.0
Temperature (water): 15° C.
Number
Percenl
Antheridial
3
Cystocarpic
0
o
•
o
Tetrasporic
9
14.7
Total
61
CD
O
•
49
Undetermined
4.9
31
TABLE VII, CONTINUED
COLLECTIONS OF GELIDIUM CARTILAGINEUM TABULATED
TO SHOW THE NUMBER AND PERCENTAGE FREQUENCY OF
THE VARIOUS TYPES OF INDIVIDUALS
Collection of April 1, 1939, Laguna Beach, California
Tide: 1:18 P.M.
-0.6
Temperature (water): 16.5° C.
Number
Percent
Antheridial
8
7.6
Cystocarpic
14
13.3
Tetrasporic
23
21.9
Undetermined
60
57.1
Total
105
32
TABLE VIII
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of February 13, 1938, Laguna Beach, California
Specimen
number
Degree of
injur y*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
1
2
1
2
1
1
3
1
2
1
1
1
1
3
3
2
2
2
fair
fair
fair
fair
fair
fair
no
fair
fair
fair
no
fair
fair
no
no
fair
fair
fair
^Explanation:
1 means slight injury
2 means pronounced injury
3 means no injury
33
TABUS VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of March 13, 1938, Point Firmin, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
3
1
3
2
1
3
2
1
2
3
2
no
fair
good
fair
fair
no
fair
good
fair
good
no
*See footnote, Page 32
54
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of April 29, 1938, Laguna Beach, California
Specimen
number
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
Degree of
injury*
2
1
1
3
1
2
2
2
2
2
2
2
2
2
1
2
2
1
2
2
2
2
2
2
2
2
2
2
1
2
1
2
2
1
2
2
1
1
Regeneration
no
no
fair
no
no
no
fair
fair
no
fair
no
fair
fair
no
no
no
no
no
fair
good
good
no
no
fair
no
good
fair
fair
no
no
no
fair
fair
fair
no
no
fair
no
*See footnote,
Page 32.
55
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of May 8, 1938, Laguna Beach, California
Specimen
number
1
2
3
4
5
6
7
8
9
10
11
12
15
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
Degree of
injury*
2
2
3
2
1
2
1
3
3
2
2
2
1
3
2
1
2
1
3
3
1
3
3
1
2
3
3
3
1
1
1
2
3
2
1
2
Regeneration
fair
fair
no
fair
good
fair
fair
no
no
fair
fair
fair
fair
no
fair
no
fair
fair
fair
fair
fair
fair
no
fair
fair
no
no
no
fair
no
fair
fair
fair
fair
good
fair
36
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of May 8, 1938, continued
Specimen
number
Degree of
injury*
Regeneration
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
1
3
3
1
1
2
2
1
2
3
2
3
3
1
1
2
1
2
2
fair
no
no
fair
fair
fair
fair
fair
fair
no
fair
no
no
no
no
fair
fair
fair
no
*See footnote, Page 32*
37
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of July 3, 1938, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
3
3
3
1
3
3
3
1
3
1
1
1
2
1
2
1
2
2
2
3
1
2
1
2
2
1
no
no
no
no
no
no
good
no
no
no
no
no
no
no
fair
no
fair
fair
fair
no
no
no
fair
fair
fair
no
*See footnote, Page 32
58
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of July 30, 1938, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
2
3
3
no
no
no
fair
no
no
no
fair
no
no
no
fair
fair
no
no
no
no
no
no
no
fair
no
no
10
11
12
13
14
15
16
17
18
19
20
21
22
23
*See footnote, Page 32
1
3
1
3
2
3
2
2
2
2
2
1
2
1
1
3
3
1
3
3
59
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection or September 25, 1938, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
2
2
2
2
1
2
2
1
2
2
1
1
1
1
3
1
1
1
2
3
3
1
3
no
no
fair
fair
fair
fair
no
no
no
no
no
no
fair
no
no
fair
no
no
good
fair
no
no
no
*See footnote, Page 32
40
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of October 23, 1938, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
2
2
2
2
2
2
2
2
1
2
2
3
1
1
1
2
2
2
2
1
2
2
2
2
2
2
2
3
1
1
1
2
2
2
1
1
no
good
fair
no
no
no
fair
fair
no
no
fair
no
no
no
good
no
fair
fair
fair
no
no
fair
fair
no
no
no
fair
no
fair
no
fair
fair
no
no
no
no
-*See footnote, Page 32
41
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of November 20, 1938, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
1
3
2
1
2
2
2
2
2
2
2
1
2
1
2
1
2
1
1
1
3
2
1
2
2
1
1
2
2
1
2
1
2
1
no
no
good
fair
fair
fair
no
no
fair
fair
fair
no
fair
fair
good
no
fair
fair
no
no
no
fair
no
no
no
no
no
fair
no
no
fair
fair
fair
good
42
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHED AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of November 20, 1958, continued
Specimen
number
Degree of
injury*
Regeneration
35
36
37
38
39
40
41
42
43
AXAX
*
45
2
fair
no
no
fair
no
fair
no
fair
no
no
fair
*See footnote, Page 32
1
3
1
3
2
2
2
2
3
1
43
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of February 2, 1939, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
12
13
14
3
3
2
2
1
3
3
2
2
2
1
2
1
1
no
no
fair
no
no
no
no
fair
no
good
no
no
no
no
*See footnote, Page 32
44
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of March 5, 1959, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
8
5
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
2
2
2
2
2
1
2
2
2
2
2
2
1
2
2
2
2
2
2
2
1
2
2
1
1
2
2
1
2
2
2
2
2
1
fair
fair
fair
good
fair
no
no
fair
no
fair
no
no
no
no
no
no
fair
no
fair
fair
no
no
fair
no
fair
fair
fair
no
fair
fair
fair
no
no
no
45
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of March 5, 1939, continued
Specimen
number
Degree of
injury*
Regeneration
35
36
37
38
39
40
1
2
1
2
1
1
no
no
no
good
no
fair
*See footnote, Page 32
46
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of April 1, 1939, Laguna Beach, California
Specimen
number
Degree of
injury*
Regeneration
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
1
2
1
1
2
2
2
1
1
2
2
2
2
2
2
2
2
2
2
2
2
1
2
2
3
2
1
2
1
3
1
2
2
2
2
fair
fair
no
fair
fair
fair
fair
fair
fair
fair
fair
no
no
no
fair
good
no
no
fair
no
no
no
good
no
no
good
fair
fair
fair
no
fair
fair
fair
fair
fair
*See footnote, Page 32*
47
TABLE VIII, CONTINUED
THE EXTENT OF INJURED BRANCHES AND THE DEGREE OF
REGENERATION AMONG SPECIMENS UNDETERMINED
AS TO TYPES OF REPRODUCTIVE ACTIVITY
Collection of April 1, 1939, continued
Specimen
number
Degree of
injury*
Regeneration
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
1
2
2
2
2
2
2
2
2
1
1
2
2
1
3
3
1
2
2
2
2
2
2
2
2
3
fair
fair
fair
fair
fair
fair
fair
fair
fair
no
no
fair
fair
no
no
no
no
fair
fair
good
no
good
no
fair
no
no
-*See footnote, Page 32.
48
TABLE IX
DEGREE OF ACTIVITY AMONG ADULT SHOOTS (x), AND
THE OCCURRENCE OF NEW BASAL SHOOTS (xj
Material from Laguna Beach, California, except for March
13, 1938 (from Point Firmin, California).
February 13, 1938
No* Good Fair
1
2
3
4
5
Slight
April 10, 1938
No.
Good Fair
x
X
X
3
4
5
6
X
6
7
8
9
X
10
7
1
2
3
4
5
6
7
8
9
10
8
x
x
9
X
X
X
X
X
X
X
10
March 13, 1938
No. Good Fair
Slight
Slight
x
x
x
x
x
x
x
x
May 8, 1938
No. Good
1
2
3
4
5
6
7
8
x
x
9
10
X
Fair
Slight
x
x
X
X
X
X
X
X
X
49
TABLE IX, CONTINUED
DEGREE OF ACTIVITY AMONG ADULT SHOOTS (x), AND
THE OCCURRENCE OF NEW BASAL SHOOTS (x)
Material from Laguna Beach, California, except for March
13, 1938 (from Point Firmin, California)
July 3, 1938
Ho. Good Fair
1
2
3
4
5
6
7
8
9
10
X
X
X
X
X
X
X
X
X
X
July 30, 1938
Ho. Good Fair
1
2
3
4
5
6
7
8
9
10
Slight
Slight
X
X
X
X
X
X
X
X
X
X
September 25, 1938
No.
Good Fair Slight
1
2
3
4
5
6
7
8
9
10
X
X
X
X
X
X
X
X
X
X
October 23, 1938
No. Good Fair
1
2
3
4
5
6
7
8
9
10
X
X
X
X
X
X
X
X
X
Slight
50
TABLE IX, CONTINUED
DEGREE OF ACTIVITY AMONG ADULT SHOOTS (x), AND
THE OCCURRENCE OF NEW BASAL SHOOTS (xj
Material from Laguna Beach, California, except for March
13, 1938 (from Point Firmin, California)
November 20, 1938
No. Good Fair
X
1
2
3
4
5
6
7
8
9
X
X
x
X
x
x
X
X
February 2, 1939
No. Good Fair
£
x
x
x
x
x
x
x
March 5, 1939
No. Good Fair
X
1
X
10
1
2
3
4
5
6
7
8
9
10
Slight
Slight
x
x
2
3
4
5
6
7
8
9
X
X
X
X
X
10
X
X
X
X
April 1, 1939
No. Good Fair
1
2
3
x
x
4
5
6
7
8
9
10
Slight
£
x
x
x
x
x
x
x
51
TABLE X
THE NUMERICAL RELATION OF FREQUENCY OF ADULT SHOOTS
(NUMERATOR) TO NEW SHOOTS (DENOMINATOR) AMONG
INDIVIDUAL HERBARIUM SPECIMENS
Material from Laguna Beach, California, except March
13, 1938 (from Point Firmin, California)
February 13, 1938
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
9/8
10/4
22/7
12/7
5/13
8/3
8/9
18/8
8/15
17/13
April 10, 1938
1
.
2.
3.
4.
5.
6.
7.
8.
9.
10.
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
6/3
5/1
6/8
21/5
6/2
4/4
11/1
6/3
9/2
4/4
78/33
12/4
4/3
5/5
8/3
3/6
6/1
4/1
10/4
10/5
1.
2.
3.
4.
5.
6.
7.
8
.
9.
10.
May 8, 1938
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
16/7
6/1
20/7
4/2
4/5
5/2
6/2
6/1
3/1
2/4
39/32
4/3
5/2
1/2
2/3
16/6
17/9
16/4
21/6
7/8
14/8
103/51
74/42
117/87
March 13, 1938
12/10
July 3, 1938
July 30, 1938
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
6/6
5/11
12/9
2/3
2/3
5/4
6/3
1/2
4/4
1/3
44/48
52
TABLE X, CONTINUED
THE NUMERICAL RELATION OF FREQUENCY OF ADULT SHOOTS
(NUMERATOR) TO NEW SHOOTS (DENOMINATOR) AMONG
INDIVIDUAL HERBARIUM SPECIMENS
Material from Laguna Beach, California except March
15, 1938 (from Point Firmin, California)
September 25, 1938
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
15/12
6/11
9/7
9/8
13/6
4/5
i/10
14/16
9/11
6/7
November 20, 1938
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
23/12
11/9
11/14
3/3
7/9
15/14
20/6
4/7
12/3
15/5
121/82
1.
2.
3.
4.
5.
6.
7.
8.
9.
10.
87/65
94/93
October 23, 1938
6/7
10/5
8/6
6/8
6/3
7/5
9/6
7/10
7/12
21/3
March 5, 1939
February 2, 1939
1
.
2.
3.
4.
5.
6.
7.
8.
9.
10.
12/2
9/3
18/3
7/3
9/1
3/10
8/3
3/3
6/6
3/9
78/43
15/6
6/8
19/6
5/5
19/3
14/6
24/13
10/9
5/2
5/12
122/70
April 1, 1939
1
.
2.
3.
4.
5.
6.
7.
8.
9.
10.
26/4
5/6
4/3
6/6
17/3
9/4
12/4
6/9
4/5
10/7
99/51
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